The first detailed history
of the evolution of the
five races of man— a pio-
neer work, a milestone of
scientific thought. It ad-
vances Darwinian theory
beyond the origin of spe-
cies to the myth-ridden
question of the origin of
subspecies, or races.
Fhe origin of races
CARLETON S. COON
AUTHOR OF THE STORY OF MAN
<»
0.0. R
A.A.K
Hitherto, it has generally been held
that the five races of man became differ-
entiated only very recently, in the last
few tens of thousands of years, after the
appearance of Homo sapiens. But Dr.
Coon, in the course of his researches,
uncovered startling evidence indicating
that, in fact, they had separated far down
on the time scale, long before Homo sa-
piens appeared and at least as early as the
time of the first 43S3ESS^ Homo erectus.
Was it possible that races, like the species to
which they belong, were capable of evolution?
If so, much of the evolution of the different
existing races may have taken place separately
and in parallel fashion over a period of hun-
dreds, rather than tens, of thousands of years.
Dr. Coon began to collect every scrap of exist-
ing information about fossil man, to find out
how many racial lines could be traced back to
the earliest evidence of man’s existence. He
ended with five lines of descent, each as old
as man himself. Now, the possibility that races
could be older than species had to be inves-
tigated. In order to reconstruct man’s ancestral
journey, Dr. Coon undertook no less than a vast
scientific exploration in time and space.
What forces, he had to know, exerted pres-
sure on that plastic primate, man, to make him
evolve from a lesser to a more sapient state? To
answer this most fundamental and exciting
question of all, Dr. Coon drew on the resources
of zoogeography, primate behavior, physiology,
and social anthropology; he surveyed the rules
of the formation of species, of the composition
of populations, of systems of mating, and of
geographical adaptation at different ecological
levels; he delved into the records of paleontol-
ogy and surveyed the relics and artifacts of a
hundred millennia. With this marshaling of
(continued on hack flap)
Illustrated with photographs, drawings,
charts, tables, and 13 maps drawn by
Rafael Palacios
JACKET DESIGN BY GEORGE GIUST
ALSO BY
C ARLETON S. COON
THE STORY OF MAN
( i954> 1962)
THE SEVEN CAVES
(i957)
These are Borzoi Books
published by Alfred A. Knopf in New York
THE ORIGIN OF RACES
THE
ORIGIN
OF
RACES
by CARLETON S. COON
i 9
6 2
NEW YORK : A L F R E D • A • K N O P F
L. C. catalog card number: 62—i4y6l
THIS IS A BORZOI BOOK,
PUBLISHED BY ALFRED A. KNOPF, INC.
Copyright © 1962 by Carleton S. Coon. All rights re-
served. No part of this book may be reproduced in any
form without permission in writing from the publisher,
except by a reviewer, who may quote brief passages and
reproduce not more than three illustrations in a review to
be printed in a magazine or newspaper. Manufactured in
the United States of America, and distributed by Ran-
dom House, Inc. Published simultaneously in Toronto,
Canada, by Random House of Canada, Limited.
FIRST EDITION
T O
FRANZ WEIDENR EICH
IN MEMORIAM
INTRODUCTION
I N 1933 I was invited to rewrite Professor W. Z. Ripley’s classic
The Races of Europe (New York: Appleton & Co.; 1899). My
completely new version of the book was published by The Mac-
millan Company in 1939. At that time I decided eventually to
write a Races of the World. For twenty years, in peace and war,
at home and on expeditions, I collected material with this task in
mind. Finally, in 1956, thanks to an Air Force contract, I was able
to make a seven months’ trip around the world, visiting countries
I had never before seen and conferring with fellow physical an-
thropologists on the way. From the end of that trip to the present
I have been engaged almost exclusively in the preparation of the
book at hand.
But this book is only half of what I set out to write. By 1959 it
was clear to me that I must write two books, one on the living, as
originally planned, and an introductory one on the ancestry of the
living races of man. By then I could see that the visible and in-
visible differences between living races could be explained only
in terms of history. Each major race had followed a pathway of
its own through the labyrinth of time. Each had been molded in
a different fashion to meet the needs of different environments,
and each had reached its own level on the evolutionary scale.
What became the first book, the one presented here, may turn
out to have been the harder to write, or so it seems now that I
have finished it and before I have allowed myself to become im-
mersed in the other. It was difficult because I had spent less time
on fossil men than on the living. Also, in 1959 I decided that the
framework for the study of fossil man should be built in two di-
Introduction
viii
mensions, time and space. Most other writers had stressed only
time, and had ignored or neglected geography.
A notable exception was Franz Weidenreich. While I was writ-
ing The Races of Europe in Cambridge, Massachusetts, he was
busy in New York, studying the Sinanthropus remains. At that
time he concluded that the peculiarities that made Sinanthropus
distinct from other fossil men were of two kinds, evolutionary and
racial. From the evolutionary point of view, Sinanthropus was
more primitive than any known living population. Racially he was
Mongoloid.
Like other premature comets of science, Weidenreich’s idea
flashed across the sky and was gone, obscured by the clouds of
incredulity released by his fellow scientists. Most of them be-
lieved, as many still do, that the living races of man could have
become differentiated from a common ancestor only after the
stage of Homo sapiens had been reached. Because Homo sapiens
was believed to have first appeared only 30,000 years ago, in the
guise of Cro-Magnon man, the living races could be only that old.
Sinanthropus was not Homo sapiens. Therefore he could not have
belonged to a modern race, the Mongoloid. Q.E.D. Or so the in-
credulous thought.
To me there was something very pat, dogmatic, and wrong
about the anti- Weidenreich point of view. For years I mulled it
over in my mind, and then I decided to collect every scrap of
existing information about every single fossil-man bone and tooth
in the world. Once I had acquired as much information as I could,
I concentrated on the dimension of space and tried to see how
many racial lines, including the Mongoloid, could be traced back
to the first instance that any kind of man had appeared on the
earth. In the end I succeeded in tracing back five, each as old as
man himself.
Realizing the enormity of my discovery in terms of its diver-
gence from accepted dogma, I knew that I must provide a theo-
retical foundation for the facts I had unearthed. The possibility
that races can be older than species had to be explored. I soon
found, by reading and through conversations with Mayr, Simpson,
and other biologists, that what I had thought a revolutionary con-
cept was so common an event in nature that others rarely both-
Introduction
ix
ered to mention it; to wit, that a species which is divided into
geographical races can evolve into a daughter species while re-
taining the same geographical races.
With this matter settled more easily than I had expected, I
needed to know what forces exerted pressure on that plastic pri-
mate, man, to make him evolve from a lesser to a more sapient
state. To satisfy this need, I delved into zoogeography, primate
behavior, physiology, and social anthropology. At the same time I
kept in touch with physiologists studying the mechanisms of ad-
aptation to heat, cold, and altitude, and went with some of them
on a field trip to southern Chile.
Because my study made it apparent that the human races had
evolved in parallel fashion, I made a brief excursion into the his-
tory, anatomy, and physiology of primates, and found many strik-
ing examples to back my theory. Meanwhile, the exciting new
discoveries regarding fossil apes and Australopithecines drew the
prehuman relatives of man forward in time past the very date of
the earliest human skull, closing a temporal if not an evolutionary
gap. These discoveries opened the possibility that the races of
man are even older than the known specimens of Homo, a possi-
bility that remains unexplored.
In the introduction to The Races of Europe I stated that I
would avoid discussion of two subjects, blood groups and racial
differences in intelligence. W. C. Boyd was about to publish his
massive compilation of blood groups.1 And I knew next to nothing
about racial intelligence and could not see that it would be very
useful when applied to regional populations of a single major
race, the Caucasoid.
The sequel to The Origin of Races promises to be full of talk
about blood and brains, but in this present book I have little to
say about these subjects — for different reasons than in 1939. De-
spite claims to the contrary, the blood groups of fossil bones can-
not be determined. Nor can dead men take intelligence tests.
However, it is a fair inference that fossil men now extinct were
less gifted than their descendants who have larger brains, that the
subspecies which crossed the evolutionary threshold into the cate-
1 W. C. Boyd: “Blood Groups,” Tabulae Biologicae , Vol. 17 (1939), pp. 113-
240.
X
Introduction
gory of Homo sapiens the earliest have evolved the most, and
that the obvious correlation between the length of time a sub-
species has been in the sapiens state and the levels of civilization
attained by some of its populations may be related phenomena.
Yet every major race, however advanced in civilization some of
its component populations have become, also contains remnant
bands of simple hunters and gatherers to remind us whence we
all came. The monkey-hunters of the forested slopes of Central
India are as Caucasoid as Charles de Gaulle, and the Ghosts of the
Yellow Leaves, who haunt the hillsides of Upper Siam and Laos,
as Mongoloid as the Mikado.
These, however, are not the main points of the book. This is a
work of history, the history of a primate genus, and in it science is
only a set of tools used to discover the pathways of human evolu-
tion— pathways that have led us from a time of obscurity to a mo-
ment of bright sunlight, with no man knows what fate lying ahead.
Carleton S. Coon
Devon, Pennsylvania
January 23, ig62
ACKNOWLEDGMENTS
In the compilation and preparation of data for this
volume I have received financial aid from one private and two gov-
ernmental institutions, as follows. In 1955 the Wenner-Gren Foun-
dation paid my way to and from the Third Panafrican Congress at
Livingstone, Northern Rhodesia, where Mrs. Coon and I were
the guests of the Rhodesian government for several weeks. Since
then the Wenner-Gren Foundation has given me two other grants.
In 1956-7 we went around the world on Contract AF33(6i6)-
6306 with ADTIC ( Arctic-Desert-Tropics-Information Center)
of the Montgomery, Alabama, U.S. Air Force Rase. Thanks are ex-
tended to Dr. Paul H. Nesbitt, chief of the ADTIC, and his staff
for their helpful suggestions and courtesies. In 1957 I received a
two-year grant from the National Science Foundation (NSF
03921). In 1959 I went to Wellington Island, Chile, as a member
of an expedition led by T. H. Hammel of the University of Penn-
sylvania Medical School and financed by the Riomedical Labora-
tory of the U.S. Air Force at the Wright-Patterson Air Force Rase,
Ohio.
I am also deeply indebted to those who make decisions in the
University of Pennsylvania, and particularly to Froelich G.
Rainey, Director of the University Museum, to Alfred Kidder II,
Associate Director, and to the Museum’s Board of Trustees,
headed by Percy C. Madeira, Jr., for allowing me ample writing
time, granting me free use of the Museum’s facilities, and, even
more important, giving me encouragement in the pursuit of what
must have been, from the point of view of the Museum, a mate-
rially unrewarding subject. I hope that this book will justify their
confidence.
To many persons we owe debts of gratitude for fine hospitality
Acknowledgments
xii
on our travels, particularly to Dr. and Mrs. Neil Ransford, M.D.,
in Bulawayo, to the Gordon Brownes in Siam, to the Gordon
Bowles in Japan, to Arthur Prager and the Robert Lindquists in
Formosa, to the late B. S. Guha in India, to the Arthur Gardiners
in Pakistan, to Colonel and Mrs. W. A. Eddy and Aramco, Saudi
Arabia, to the late Baron and the Baroness A. C. Blanc in Rome,
to Lidio Cipriani in Florence, to M. and Mine Frangois Trives in
Paris, and to our son C. S. Coon, Jr., and his wife and children, in
many places.
My wife, Lisa Dougherty Coon, went with me everywhere
mentioned except Chile, and where she went she kept me in good
health. She also created all but six and a half of the line drawings
in this book.
No one can write a book of this scope without friends. Reprint
swapping, informal correspondence, and conversations at aca-
demic meetings and elsewhere are as important as library research,
which itself is greatly facilitated by the interest and good will of
librarians.
Below are listed the names, without rank or title, of some of
those who have helped me, in one way or other, in terms of
principal categories of assistance. Most heartily I thank them one
and all.
FIELD TRIPS
Gordon T. Bowles
A. C. Blanc f
J. Desmond Clark
B. S. Guha f
H. T. Hamm el
Louis Leakey
HRH Peter
Roger Summers
Tokyo, Johns Hopkins
Rome
Livingstone, Berkeley
Ranchi
U. of Pennsylvania, Yale
Nairobi
Prince of Greece and Denmark
Bulawayo
TAXONOMY
F. Clark Howell
Wm. W. Howells
Ernst Mayr
Chicago
Harvard
Harvard
Acknowledgments xiii
Brian Patterson
Alfred S. Romer
George G. Simpson
Wm. L. Straus, Jr.
CHROMOSOMES
M. A. Bender
John Buettner-Janusch
E. H. Y. Chu
David Hungerford
CHRONOLOGY
Bruce Howe
G. H. R. von Koenigswald
Charles E. Stearns
Elisabeth K. Ralph
Kenneth P. Oakley
SKELETAL MATERIAL AND CASTS
Don Brothwell
Georgi Debetz
Paul Deraniyagala
A. C. Hoffman
S. Kodama
Louis Leakey
P. N. Mitra
Emily Pettinos
Ronald Singer
John T. Robinson
Wm. L. Straus, Jr.
H. Suzuki
Philip T. Tobias
Harvard
Harvard
Harvard
Johns Hopkins
Oak Ridge
Yale
Oak Ridge
Institute for Cancer Research,
Fox Chase, Philadelphia
Harvard
Utrecht
Tufts
University of Pennsylvania
British Museum (Nat. Hist.)
British Museum (Nat. Hist.)
Academy of Science, Moscow
Colombo Museum
Bloemfontein
Sapporo
Nairobi
Calcutta
University Museum, Philadel-
phia
Capetown, Chicago
Pretoria
Johns Hopkins
Tokyo
Johannesburg
XIV
Acknowledgments
GENERAL HELP, PARTICULARLY IN THE SUMMER OF 1958
Edward E. Hunt, Jr. Harvard and Forsythe
( Boston)
MAKING FLESH RECONSTRUCTIONS OF FOSSIL MEN
Maurice P. Coon Cambridge, Massachusetts
INFORMATION ABOUT THE TIWI,
Jane C. Goodale
RESEARCH: LIBRARIANS
Margaret Currier
Cynthia Griffin
Margaret Palmer
BIBLIOGRAPHY
Janet M. Kliment
GLOSSARY SELECTION
Mary S. Huhn
WORK ON PHOTOGRAPHS
David Crownover
Caroline Dosker
Jane C. Goodale
Doris Nicholas
chapter 3
University Museum,
Bryn Mawr
Peabody Museum, Harvard
University Museum, University
of Pennsylvania
Dental School, University of
Pennsylvania
J
University Museum, University
of Pennsylvania
Devon, Pennsylvania
University Museum
University Museum
University Museum,
Bryn Mawr
University Museum
Acknowledgments xv
But this is not all. Books need publishers as well as authors.
I am deeply indebted to Alfred A. Knopf for the privilege of
having had Harold Strauss as editor of this volume, with the
valuable assistance of Howard Fertig, Sophie Wilkins, and Carmen
Gomezplata, and for splendid treatment at the hands of William
Koshland.
I am also very happy that my British publisher, Jonanthan Cape
Ltd., which has stood by me for thirty years, will publish this book
in England.
CONTENTS
1. THE PROBLEM OF RACIAL ORIGINS
ON THE ANTIQUITY OF RACES; THE PROBLEMS OF HUMAN
TAXONOMY: THE GENUS; HOMO SAPIENS; THE SPECIES
CONCEPT; THE SPATIAL REQUIREMENTS OF SPECIES AND
THEIR GEOGRAPHICAL DIFFERENTIATION; THE SUBSPECIES;
MOSAICS, CLINES, LOCAL RACES, AND RACIAL TYPES; THE
DIFFERENTIATION OF SPECIES; BALANCED POLYMORPHISM;
ON THE TIMING OF THE INDIVIDUAL GROWTH CYCLE; ON
SIZE AND FORM: ALLOMETRY; ON SEXUAL DIMORPHISM;
HOW SPECIES HAVE EVOLVED; ON THE LIFE SPANS OF
MAMMALIAN SPECIES; GENETIC PRINCIPLES AND THE
ORIGINS OF RACES
2. EVOLUTION THROUGH ENVIRONMENTAL
ADAPTATION
BODY SIZE, FOOD, SPACE, AND CLIMATE; THE FACE OF THE
EARTH; LAND MASSES; BARRIERS AND BREEDING AREAS; GE-
NETIC DRIFT; THE DOMINANCE OF GROUPS; THE SIX FAU-
NAL REGIONS; WALLACEA; THE FAUNAL REGIONS AND
HUMAN ORIGINS AND MOVEMENTS; ENVIRONMENTAL
ADAPTATION AND EARLY MAN; THE RULES OF BERGMANN
AND ALLEN; NOSE FORM AND CLIMATE; PHYSIOLOGICAL
ADAPTATION TO COLD; HEAT ADAPTATION; THE SIGNIFI-
CANCE OF ADAPTATION TO HEAT AND COLD; ADAPTATION
TO ALTITUDE
3. EVOLUTION THROUGH SOCIAL
ADAPTATION
LEADERSHIP, COMMUNICATION, AND BRAIN GROWTH; ON
THE ANTIQUITY OF A HUMAN TYPE OF SOCIETY: THE BE-
GINNING OF HUNTING; THE MATING SYSTEMS OF OTHER
xviii Contents
ANIMALS; THE SEXUAL BEHAVIOR OF PRIMATES, INCLUD-
ING HOMO SAPIENS; THE BEGINNINGS OF HUMAN SOCIETY;
SEXUAL SELECTION AMONG HIGHER PRIMATES; SPEECH,
HUNTING, AND SOCIAL STRUCTURE; RITUAL, LANGUAGE,
AND THE RITES OF PASSAGE; THE DISCOVERY OF FIRE AND
THE CONVERSION OF ENERGY INTO SOCIAL STRUCTURE;
THE EVIDENCE OF LIVING FOOD-GATHERING SOCIETIES: THE
AUSTRALIAN ABORIGINES; THE ARCHAIC SOCIETY OF THE
TIWI; ON COMPARING THE CULTURES OF LIVING FOOD
GATHERERS AND THOSE OF FOSSIL MEN; POPULATION SIZE
AMONG FOOD GATHERERS; SYSTEMS OF MATING AMONG
FOOD GATHERERS; THE LONGEVITY OF FOSSIL MEN; THE
ROLE OF ISOLATING MECHANISMS IN HUMAN EVOLUTION;
ADAPTATION TO CROWDING: A NEW THEORY OF EVOLU-
TION BY SUCCESSION; DWARFING AS A SOLUTION TO THE
PROBLEM OF CROWDING; THE ENDOCRINES AND TEMPERA-
MENT; PARALLELS BETWEEN ANIMAL DOMESTICATION
AND SOCIAL ADAPTATION; THE UNIQUE ADAPTATIONS OF
THE GENUS HOMO
4. THE ORDER OF PRIMATES
PRIMATE STUDIES AND THE CLASSIFICATION OF HUMAN
RACES; THE CLASSIFICATION OF PRIMATES; THE PROSIM-
IANS; THE TREE SHREWS; THE LEMURS; THE LORISES;
THE TARSIERS; THE LIVING PLATYRRHINES : THE SOUTH
AMERICAN MONKEYS; THE LIVING CERCOPITHECIDAE : OLD
WORLD MONKEYS; THE LEAF-EATING COLOBINAE; THE
CERCOPITHECINAE; THE ANTHROPOID APES; THE GIBBONS:
SYMPHALANGUS AND HYLOBATES; THE ORANGUTAN
(PONGO); THE CHIMPANZEE (pan); THE GORILLA ( GO-
RILLA); THE HOMINIDAE ( HOMO )
5. MAN’S PLACE AMONG THE PRIMATES
THE BEARING OF PRIMATE STUDIES ON RACIAL ORIGINS; TO
BRACHIATE OR NOT TO BRACHIATE; THE BEARING OF
HOMINID TEETH ON THE ERECT POSTURE; A FEW DETAILS
OF THE POSTURE STORY; THE EVIDENCE OF TEETH; THE
EVIDENCE OF EMBRYOLOGY; DIFFERENCES IN POSTNATAL
GROWTH; PHYSIOLOGICAL CLUES TO OUR RELATIONSHIPS
WITH OTHER PRIMATES; PARASITES AND PRIMATES; THE
Contents
xix
COMP ARISON OF PRIMATE CHROMOSOMES; THE EVIDENCE
OF BEHAVIOR
6. THE FOSSIL RECORD: FROM LEMURS TO
SWAMP APES 186
ON THE SCARCITY OF PRIMATE FOSSILS; THE PRIMATE
TIME SCALE; PRIMATE PALEONTOLOGY AS A WHOLE; THE
PROSIMIAN PROLIFERATION; THE EVOLUTION OF THE PLAT-
YRRHINES; THE EVOLUTION OF THE CATARRHINES; THE
GIBBON LINE; THE ANCESTORS OF THE THREE LIVING GREAT
APES; PROCONSUL; DRYOPITHECUS IN EUROPE AND ASIA;
RAMAPITHECUS, A POSSIBLE ANCESTOR OF THE HOMINIDS;
THE FORT TERNAN PRIMATE; THE PLEISTOCENE APES OF
CHINA; POSSIBLE SURVIVALS OF CHINESE APES; HOMINOIDS
AND HOMINIDS; OREOPITHECUS BAMBOLII, THE SWAMP
APE; FOSSIL PRIMATES AND HUMAN EVOLUTION
7. THE EARLIEST HOMINIDS 217
THE ORIGIN OF THE HOMINIDS; AUSTRALOPITHECUS AND
HOMO; THE LOWER PLEISTOCENE; THE NEW DATING FOR
THE LOWER PLEISTOCENE; THE EVIDENCE OF TOOLS AND
FIRE IN THE LOWER PLEISTOCENE; GEOGRAPHY AND NUM-
BERS OF EARLY HOMINIDS; THE SOUTH AFRICAN AUSTRA-
LOPITHECINES : TIME, SPACE, AND TAXONOMY; THE
AUSTRALOPITHECINE CAVE SITES; DID THE AUSTRALOPITHE-
CINES MAKE TOOLS?; THE POSTCRANIAL SKELETONS OF
THE SOUTH AFRICAN AUSTRALOPITHECINES; THE STERK-
FONTEIN VERTEBRAE AND RIBS; THE PELVIS OF AUSTRALO-
PITHECUS; THE LEGS AND FEET OF AUSTRALOPITHECUS;
THE SHOULDER GIRDLE OF AUSTRALOPITHECUS; THE ARMS
AND HANDS OF AUSTRALOPITHECUS; AUSTRALOPITHECUS, A
PRIMATE MERMAID OR A UNIQUE HOMINID?; THE SKULLS,
JAWS, AND TEETH OF AUSTRALOPITHECUS; THE BRAIN
CASE AND BRAIN OF AUSTRALOPITHECUS; THE FACES OF
THE AUSTRALOPITHECINES; THE AUSTRALOPITHECINE
JAWS; THE TEETH OF AUSTRALOPITHECUS; THE EARLY
HOMINIDS FROM EAST AFRICA; THE OLDUVAI CHILD; THE
CHILD’S MANDIBLE; THE CHILD’S TEETH; THE CHILD’S
PARIETAL BONES; THE FOOT ACCOMPANYING THE CHILD’S
REMAINS; THE COLLARBONE, HAND, AND FINGERS; THE
EVOLUTIONARY AND TAXONOMIC POSITION OF THE OLDU-
XX
Contents
VAI CHILD; ZINJANTHROPUS: HIS TOOLS, DIET, AND ACTIV-
ITIES; THE ANATOMY OF ZINJANTHROPUS: HIS CRANIUM;
THE TEETH OF ZINJANTHROPUS; THE LEG BONES ATTRI-
BUTED TO ZINJANTHROPUS; THE STATUS OF ZINJANTHRO-
PUS; THE SPECIMEN FROM LAKE EYASI, TANGANYIKA; THE
KANAM MANDIBLE; THE AUSTRALOPITHECINE FROM THE
REPUBLIC OF TCHAD; THE FOSSIL HOMINID OF TELL UBEI-
DIYA, JORDAN VALLEY; THE MEGANTHROPUS MANDIBLES
FROM JAVA; THE DRUGSTORE AUSTRALOPITHECINES OF
CHINA; THE REPLACEMENT OF AUSTRALOPITHECUS BY
HOMO
8. AN INTRODUCTION TO FOSSIL MAN
OF TIME, SPACE, GRADES, AND LINES; THE DIMENSION OF
TIME; THE DIMENSION OF SPACE: GLACIAL GEOGRAPHY;
THE TEMPORAL AND SPATIAL DISTRIBUTION OF FOSSIL
MAN SITES; TIME, SPACE, AND PALEOLITHIC TOOLS; THE
CHRONOLOGY AND DISTRIBUTION OF THE USE OF FIRE;
GRADES AND SPECIES OF FOSSIL MEN; THE SAPIENS-
ERECTUS THRESHOLD: THE EVIDENCE OF BRAIN SIZE; THE
EVIDENCE OF CRANIAL FORM; THE EVIDENCE OF TOOTH
SIZE; A BRAIN-SIZE TO TOOTH-SIZE INDEX; EVOLUTIONARY
CHANGES WITHIN HOMO SAPIENS: THE RISE OF THE CHIN;
LINES AND SUBSPECIES OF FOSSIL MEN: THE EVIDENCE OF
TEETH; RACIAL VARIATIONS IN THE FORM AND STRUCTURE
OF TEETH; FACIAL FLATNESS AS A CRITERION OF RACE;
RACIAL ORIGINS AND RACIAL CONTINUITIES
9. PITHECANTHROPUS AND
THE AUSTRALOIDS
THE PITHECANTHROPUS LINE; THE PITHECANTHROPUS-
AU STRALOID SKELETAL MATERIAL; FOSSIL MEN FROM THE
D JETTS BEDS OF JAVA; PITHECANTHROPUS 4; THE PITHE-
CANTHROPUS MANDIBLES FROM THE DJETIS BED; THE
BRAIN CASE OF THE INFANT MODJOKERTENSIS; MEN OF
THE TRINIL FAUNA; THE PITHECANTHROPUS THIGHBONES;
THE TEETH OF PITHECANTHROPUS; THE THIRD KNOWN
HUMAN POPULATION OF JAVA: SOLO MAN; SEX, AGE, AND
INJURIES OF THE ELEVEN SKULLS; THE RACIAL ANATOMY
OF THE NGANDONG SKULLCAPS; THE FACE OF SOLO MAN;
THE NGANDONG LEG BONES; WHAT NAME, MR. SOLO?; THE
Contents
xxi
SOLO-LIKE BRAIN CASE FROM AITAPE, NEW GUINEA; THE
FOURTH KNOWN HUMAN POPULATION OF JAVA: WADJAK
MAN; THE WADJAK BRAIN CASES; THE WADJAK FACES;
THE WADJAK MANDIBLES; THE WADJAK DENTITION; THE
SIGNIFICANCE OF WADJAK; FOSSIL MAN IN AUSTRALIA;
THE KEILOR SKULL; THE TALGAI SKULL; THE COHUNA
SKULL; THE PITHECANTHROPUS-AUSTRALOID LINE; HUMAN
EVOLUTION NORTH OF JAVA IN THE PLEISTOCENE; THE
MAPA SKULLCAP; THE UPPER PLEISTOCENE SKULL FROM
NIAH CAVE, NORTH BORNEO; THE MESOLITHIC-NEOLITHIC
TRANSITION IN INDONESIA; MESOLITHIC AND NEOLITHIC
REMAINS FROM INDOCHINA; PREHISTORIC POPULATIONS
OF THE WESTERN ORIENTAL REGION; THE TAXONOMY OF
THE AUSTRALOID SUBSPECIES
10. SINANTHROPUS AND THE MONGOLOIDS 428
THE LIVING MONGOLOIDS AND THE SKELETONS OF THEIR
ANCESTORS; SINANTHROPUS PEKINENSIS : TIME, PLACE, AND
PEOPLE; THE TAXONOMY OF SINANTHROPUS; THE SINAN-
THROPUS BRAIN CASE; THE FACE OF SINANTHROPUS; THE
MANDIBLES OF SINANTHROPUS; THE TEETH OF SINAN-
THROPUS; THE LEG BONES OF SINANTHROPUS; THE UPPER
EXTREMITY OF SINANTHROPUS; THE POSITION OF SINAN-
THROPUS ON THE HUMAN FAMILY TREE; LATE MIDDLE
PLEISTOCENE FINDS IN CHINA AND JAPAN; THE TING-TSUN
TEETH; THE CHANGYANG MAXILLA; THE SPECIMEN FROM
MAPA, KWANGTUNG; THE HUMERUS SHAFT FROM USHI-
KAWA QUARRY, JAPAN; THE UPPER PLEISTOCENE WOMAN
FROM TZE-YANG, SZECHUAN; THE UPPER PLEISTOCENE
MAN FROM LIU-KIANG, KWANGSI; THE LIU-KIANG POST-
CRANIAL BONES; THE TOOTH OF SJARA-OSSO-GOL, ORDOS;
THE REMAINS FROM TI-SHAO-GOU-WAN, ORDOS; THE UP-
PER PLEISTOCENE REMAINS FROM CENTRAL HONSHU, JA-
PAN; THE PEOPLE OF THE UPPER CAVE AT CHOUKOUTIEN;
THE SPECIMEN FROM KAIt’o-TUNG, LEIPIN, KWANGSI;
POST-PLEISTOCENE SKELETONS; AMERICA: THE WESTERN
EXTENSION OF THE MONGOLOID REALM; CONCLUSION
XXII
Contents
11. THE CAUCASOIDS
THE CAUCASOID HOME; POSSIBLE CONTACTS BETWEEN SUB-
SPECIES AND CAUCASOID EVOLUTION; CONTINUITY AND
CHANGE IN THE CAUCASOID QUADRANT; THE MAUER MAN-
DIBLE, OR HEIDELBERG JAW; THE STEINHEIM CRANIUM;
THE SWANSCOMBE CRANIAL BONES; EUROPEAN FOSSIL
MEN OF THE EARLY UPPER PLEISTOCENE; FONTECHEVADE;
SACCO PASTORE; THE EHRINGSDORF REMAINS; THE STONE
BRAIN FROM GANOVCE, CZECHOSLOVAKIA; THE ROUND-
HEADED PEOPLE OF KRAPINA; THE MANDIBLES OF THE
EUROPEANS OF THE LAST INTERGLACIAL PERIOD; THE
TEETH OF THE EUROPEANS OF THE LAST INTERGLACIAL;
POSTCRANIAL BONES OF THE LAST INTERGLACIAL: THE
EVIDENCE FROM KRAPINA; THE “NEANDERTHALS” OF EU-
ROPE; THE NUMBERS AND DISTRIBUTION OF THE NEANDER-
THALS; THE WESTERN NEANDERTHALS; THE WESTERN
NEANDERTHAL CRANIA; THE WESTERN NEANDERTHAL
MANDIBLES; THE TEETH OF THE WESTERN NEANDERTHALS;
THE POSTCRANIAL SKELETONS OF THE WESTERN NEANDER-
THALS; THE HEIGHT AND BUILD OF THE WESTERN NEAN-
DERTHALS; THE FATE OF THE WESTERN NEANDERTHALS;
THE CENTRAL EUROPEAN NEANDERTHALS; THE THREE
MANDIBLES; THE POSTCRANIAL BONES FROM SUBALYUK;
THE SUBALYUK CHILD S SKELETON; THE RUMANIAN NEAN-
DERTHAL TOE BONE; THE SIGNIFICANCE OF THE NEANDER-
THAL REMAINS FROM CENTRAL EUROPE; NEANDERTHAL
REMAINS FROM THE SOVIET UNION; THE KIIK-KOBA TOOTH
AND LIMB BONES; THE INFANT SKELETON OF STAROSELE;
THE YOUTHFUL NEANDERTHAL OF TESHIK-TASH; THE EAST-
ERN NEANDERTHALS OF SHANIDAR; THE INHABITANTS OF
PALESTINE DURING WURM I; TABUN AND GALILEE; THE
SKHUL SKULLS: NO. 4 AND HIS GROUP; THE SKULL OF
SKHUL 5; THE MOUNT CARMEL TEETH; THE POSTCRANIAL
SKELETONS OF THE SKHUL POPULATION; THE MEANING OF
THE MOUNT CARMEL SKELETONS; EGBERT, THE BOY FROM
KSAR AKIL; MORE ABOUT NEANDERTHAL ORIGINS; THE UP-
PER PALEOLITHIC PEOPLE AND THEIR CULTURE; UPPER
PALEOLITHIC SITES IN SPACE AND TIME; THE RACIAL
CHARACTERISTICS OF THE UPPER PALEOLITHIC EUROPEANS;
Contents
XXlll
THE FATE OF THE UPPER PALEOLITHIC EUROPEANS; THEER
ASIATIC RELATIVES
12. AFRICA
THE DARKEST CONTINENT; FOSSIL MAN IN NORTH AFRICA:
THE TERNEFINE-TANGIER LINE; THE TERNEFINE DISCOVER-
IES; THE LITORINA CAVE MANDIBLE; THE MANDIBLE FROM
SMUGGLERS CAVE, TEMARA, MOROCCO; THE RABAT RE-
MAINS; TANGIER MAN; THE TAFORALT CRANIAL FRAG-
MENT; THE TERNEFINE-TANGIER LINE, CANNIBALS, AND
BUSHMEN; THE MANDIBLE FROM HAUA FTEAH, CYRE-
NAICA; THE EARLIEST CAUCASOID INVADERS OF NORTH
AFRICA: THE MOUILLIANS; THE CAPSIANS; THE RACIAL
ANATOMY OF THE MESOLITHIC NORTH AFRICANS; HUMAN
EVOLUTION IN AFRICA SOUTH OF THE SAHARA; THE “MILK”
TEETH FROM OLDUVAI; A POSSIBLE NEGRO EVOLUTIONARY
LINE; THE CHELLIAN-3 SKULL FROM OLDUVAI; THE KAN-
JERA SPECIMENS; THE SALDANHA BAY SKULLCAP; THE
BROKEN HILL OR RHODESIAN SPECIMENS; THE CRANIAL
FRAGMENTS FROM LAKE EYASI, TANGANYIKA; THE MAN-
DIBULAR FRAGMENT FROM DIRE DAWA, ETHIOPIA; THE
MANDIBLE FROM THE CAVE OF HEARTHS; THE CAPE FLATS
SKULL; THE BORDER CAVE SKULL; THE CAPSIAN SETTLERS
OF THE WHITE HIGHLANDS; THE ORIGIN OF THE CAPOIDS;
THE SINGA SKULL FROM THE SUDAN; THE HOMA SHELL-
MOUND SKULLS; THE BOSKOP BRAIN CASE AND THE “bOS-
KOP RACE ; THE FLORISBAD CRANIAL FRAGMENT; THE
FORMATION OF THE MODERN CAPOID PEOPLES; THE EAR-
LIEST SKELETONS OF MODERN NEGROES; DO THE PYGMIES
HOLD THE ANSWER?; WAS AFRICA THE CRADLE OF MAN-
KIND?
588
13. THE DEAD AND THE LIVING
657
STATISTICAL APPENDIX
BIBLIOGRAPHY
GLOSSARY
667
685
711
follows page 724
INDEX
PLATES
FOLLOWING PAGE 82
I Australoid: A Tiwi from Melville Island
Carleton S. Coon
II Mongoloid: A Formosan Aborigine
Carleton S. Coon
III Caucasoid: A Pathan
Wilfred Thesiger
IV Congoid: A Shilluk
Lidio Cipriani
V Capoid: A Bushman Woman
Film Study Center, Peabody Museum,
Harvard University
VI Environmental Adaptation: Two Dinka Girls
Lidio Cipriani
VII Dr. Kristian Lange-Andersen and Lucho
Carleton S. Coon
Bushmen of the Kalahari
N. R. Farbman, Life
VIII Leadership at a Tiwi Funeral
Jane C. Goodale
IX Prosimians: Common Lemur
Eric Kirkland, F.R.P.S.
Ring-tailed Lemur
Eric Kirkland, F.R.P.S.
Slender Loris
Fox, from Pictorial Parade
Tarsius
Philadelphia Zoo
X New World Monkeys: Marmoset
Eric Kirkland, F.R.P.S.
Capuchin
Eric Kirkland, F.R.P.S.
Ornate Spider Monkey
W. Marynowicz, F.I.B.P., F.R.P.S.
XXVI
Plates
XI Old World Monkeys: Pig-tailed Macaque
Eric Kirkland, F.R.P.S.
Red-capped Mangabey
Eric Kirkland, F.R.P.S.
Brazza Monkey
Eric Kirkland, F.R.P.S.
Patas Monkey
Eric Kirkland, F.R.P.S.
XII Old World Monkeys: Mandrill
New York Zoological Society
Proboscis Monkey
Van Nostrand, San Diego Zoo
XIII Apes: Gibbon
University Museum, University of Pennsylvania
Siamang
Van Nostrand, San Diego Zoo
XIV Orangutan
W. Marynowicz, F.I.R.P., F.R.P.S.
XV Chimpanzee
Eric Kirkland, F.R.P.S.
XVI Mountain Gorilla
W. Marynowicz, F.I.B.P., F.R.P.S.
following page 370
XVII Pygmies: A Luzon Negrito Woman
Carleton S. Coon
XVIII Pygmies: Onges from Little Andaman Island
Lidio Cipriani
XIX Pygmies: A Kadar from the Cardamon Hills
Carleton S. Coon
XX A Pygmy from the Congo
Lidio Cipriani
XXI The Karyotype of Man
David PIungerford, The Institute for Cancer Research,
Fox Chase, Philadelphia
Human Chromosomes
XXII Zinjanthropus Skull, Three Views
Des Bartlett, Armand Denis Productions, Inc.;
Courtesy National Geographic
XXIII Zinjanthropus Palate
Des Bartlett, Armand Denis Productions, Inc.;
Courtesy National Geographic
XXIV Peculiarities of Tooth Structure
a. b. c. by A. A. Dahlberg; d. e. f. by P. O. Pedersen
XXV
XXVI
XXVII
XXVIII
XXIX
XXX
XXXI
XXXII
Plates
Broken Hill, Two Views
By permission of the British Museum
(Natural History)
Saldanha Bay
Carleton S. Coon
La Ferrassie 1, Three Views
George Quay, University Museum, University
of Pennsylvania
Teshik-Tash, Three Views
George Quay, University Museum, University
of Pennsylvania
Skhul 5
George Quay, University Museum, University
of Pennsylvania
Jebel Qafza 6
H. V. Vallois, Musee de l’Homme, Paris
Grimaldi; Florisbad
George Quay, University Museum, University
of Pennsylvania
Flesh Beconstructions by Maurice P. Coon
Wayland Minot
Flesh Reconstructions by Maurice P. Coon
Wayland Minot
An Australian Aborigine and a Chinese Sage
Carleton S. Coon
xxvii
DRAWINGS
FIG.
1. How One Polytypic Species Can Evolve into Another
2. The Speech Organs of Primates
3. The Skull of an Aye- Aye
4. The Molars of Old World Monkeys and Apes
5. The Jumping Skeleton
6. Transverse Rib-Cage Sections of Jumpers,
Brachiators, and Men
7. The Carrying Angle in Apes and Men
8. Feet of Apes and Men
9. Occlusion of Canines in Apes and Hominids
10. Body Proportions of Newborn Primates
11. Changes in Skull Form from Newborn to Adult
12. The Primate Time Scale: The Cenozoic Clock
13. Mesopithecus, a Miocene Leaf -eating Monkey ( Colobinae )
14. Proconsul africanus
15. Ramapithecus brevirostis
16. The Skull of Oreopithecus: Hiirzeler’s Reconstruction
17. The Skull of Oreopithecus: Drawn from a Photograph
18. The Specialized Dentition of Oreopithecus
19. The Pelvis and Femora of Oreopithecus
20. The Anatomy of the Human Pelvic Bone ( Os coxae )
21. Pelvic Bones of Ape, Australopithecus, and Man
22. The Distal End of the Femur of Australopithecus,
Ape, and Man
23. The Bones of the Human Foot, Seen from Above
24. The Astragalus of the Australopithecines and
Other Primates
25. The Foot Bones Found with the Olduvai Child
29
75
127
135
155
157
158
160
163
167
169
188
i94
200
204
210
210
211
213
242
243
245
246
248
Drawings
xxix
26. The Australopithecine Scapula
27. The Australopithecine Hand Bones
28. Skull Profiles of Australopithecines, Gorilla, and
Baboon
29. From Proconsul to the Smaller Australopithecines
to the Larger Ones
30. Variations in the Area of Neck-Muscle Attach-
ment in Apes and Australopithecines
31- Sections Through the Nasal Passages of Australo-
pithecus, Ape, and Man
32. The Anatomy of the Mandible (Swartkrans, female)
33. The Australopithecine Mandibles
34a. Australopithecine Teeth: Incisors
34b. Australopithecine Teeth: Canines
35- Australopithecine Teeth: Premolars and Molars
36. Irregularly Shaped Molars of Australopithecus robustus
37. The Upper Canine and First Premolar in Aus-
tralopithecines, Apes, and Men
38. The Teeth of Telanthropus
39. The Australopithecine Features of the Lower
First Molar of Meganthropus
40. A Section Through the Pleistocene Beds at Olduvai
Gorge
41. The Olduvai Child’s Mandible
42. A Primate Family Tree
43. Basic Tools of Early Men
44. Grades and Lines of Fossil Hominids
45. Anatomy of the Skull
46. Sagittal Arcs and Chords in Homo erectus and
Homo sapiens
47. The Lateral Pterygoid Muscles and the Chin
48. How Brow Bidges Protect the Eyes against Blows
49. Racial Variations in Tooth Structure: Shoveling
and Ridging
50. Racial Variations in Tooth Structure: the Pre-
molar Cone
51. Racial Variations in Tooth Structure: the Cingu-
lum, Wrinkling, Taurodontism, and Ena-
mel Extensions
52. Lower Molar Crown Patterns
53. The Four Indices of Facial Flatness
249
254
257
261
262
263
265
266
269
270
274
274
275
276
279
281
303
326
335
338
342
348
350
355
357
358
361
366
XXX
Drawings
54. Transverse Sections of the Skulls of a Female Go-
rilla, Pithecanthropus 4, and Solo 11 378
55. The Faces of Homo erectus 379
56. Mandibles of Meganthropus, Pithecanthropus B,
and Wadjak 2 381
57. The skulls of Homo erectus and Homo sapiens at
the Age of Two 383
58. Profiles: Australoid Skulls from Trinil to Niah 396
59. Middle Meningeal Artery Patterns of Fossil Men 443
60. Profiles: from Sinanthropus to the Upper Cave
Male 444
61. Alveolar Prognathism in Sinanthropus and in
Modern Chinese 446
62. Mandibles: Sinanthropus and Ternefine 3 448
63. Torus mandibularis 451
64. The Continuity of Mongoloid Teeth: Shovel In-
cisors from Sinanthropus to the Recent
Chinese 454
65. Mandibles: Krapina J, Ehringsdorf, Montmaurin,
and Heidelberg 490
66. Profiles: Steinheim, Swanscombe, and
Fontechevade 493
67. Profiles: Saccopastore 1, Krapina, and
Ehringsdorf 501
68. Saccopastore Inca Bones 503
69. The Mask of Krapina 509
70. From Neanderthal to Nordic in Wiirm I 529
71. Why the Neanderthals Were Not Homo erectus:
Occipital Views of Six Skulls 530
72. Caucasoid Neanderthal Mandibles: Skhul 4,
Tabun 2, La Ferrassie 1, and Circeo 3 536
73. Mandibles of Heidelberg, La Ferrassie 1, and
Skhul 4, Seen from Above 538
74. The Upper Incisors of Neanderthals and Other
Early Caucasoids 54°
75. Neanderthal Hands and Feet 55^
76. The Human Face and Hand in Upper Paleolithic
Art 586
77. The Ternefine Parietal 592
78. Mandibles: Ternefine x and Rabat 594
79. The Tangier Maxilla and Teeth 599
Drawings xxxi
80. The Molar from Olduvai Bed II g1;i
81. Profiles: Chellian 3, Saldanha, and Broken Hill 616
82. Profiles: A Possible Negro Line— Broken Hill,
Cape Flats, and Border Cave 624
83. The Florisbad Site g
84. The Capoid Line: Profiles of Homa, Fish Hoek,
and a Modern Bushman 646
MAPS
DRAWN BY RAFAEL PALACIOS
1 The Five Subspecies of Homo sapiens in a.d. 14Q2 6-7
2 The Sunda and Sahid Shelves and Wallacea 45
3 The Six Faunal Regions of Sclater and Wallace 51
4 Distribution of Living Primates 124-125
5 Faunal Movements into Java 224
6 South African Australopithecine Sites 233
7 East African Australopithecine and Early Man Sites 280
8 The Wiirm Glaciation in Europe and Contemporary
Sea Levels g!g
9 Australoid Fossil Man Sites 372~373
10 Mongoloid Fossil Man Sites 429
11 Human Skeletal Remains in Europe, Western Asia,
and North Africa 483
12 Fossil Man Sites in Africa South of and in the Sahara 615
13 Shifts of Human Subspecies from Pleistocene to Post-
Pleistocene 659
TABLES
1 Order of Primates, Living Genera
2 Numbers of Chromosomes Among the Primates
3 The Cenozoic Era in Millions of Years
4 The Distribution of Early Hominids
5 Australopithecine Postcranial Bones
6 Skidls, Jaws, and Teeth of Australopithecines
7 Numbers of Australopithecine Teeth
8 Crown Dimensions of Australopithecine Teeth
9 A Comparison of the Crown Areas of Canines and Pre-
molars in Australopithecus and Homo
10 Tentative Cranial Measurements and Indices of the
Australopithecines and of Proconsul africanus
n Fossil-Man Sites in Time and Space
12 The Brain-Palate Index and the Brain-Molar Index
13 Flowers Index
14 Racial Variations in Tooth Form
15 The Four Indices of Facial Flatness of Woo and Morant
16 Pithecanthropus- Australoid Skeletal Material
17 The Ngandong Skulls
18 Dimensions of the Hypophyseal Fossa
19 Early Skeletal Material from China and Japan
20 The Sinanthropus Specimens by Sex and Age
21 Loci and Sex of Sinanthropus Specimens
22 Internal Dimensions of the Sinanthropus and Solo
Skulls
23 Facial Dimensions of Sinanthropus ( Female ) and
Wadjak 1
24 Angles of Inclination and Indices of Robusticity of
Sinanthropus and Other Mandibles
122
180-182
188
231
241
256
268
272
273
291
322
345
353
354
367
376
392
395
430
433
434
439
445
450
XXXIV
Tables
25 Pre-Wurm Fossil Man Remains from Europe and
Western Asia
26 Internal Dimensions and Capacities of Steinheim and
of Female erectus Skulls
27 Postcranial Bones from Krapina
28 Neanderthal and Other Remains of Wiirm I or Later
29 Simple Dimensions of the Neanderthal Cranial Base
30 Upper Paleolithic Fossil Man Sites
31 Pre-Mouillian Skeletal Material from North Africa
32 Mouillian Skeletal Material
33 Capsian Skeletal Material
34 Skeletal Material from Africa South of and Including
the Sahara
35 The Teeth from Bed II of Olduvai
APPENDIX
36 Arcs and Chords of the Frontal, Parietal, and Occipital
Bones in the Sagittal Plane
3 7 Cranial Dimensions and Indices
38 Dimensions and Indices of Mandibles
39 Dimensions and Indices of Teeth
487
494
517
524
53i
580
591
605
606
612
613
667
668
675
678
PERIODICALS
AND THEIR ABBREVIATIONS
AA
AAE
AAnz
AB
ActG
AE
AEB
Afbica
AGMG
AIPH
AJAn
AJHG
AJPA
AJSc
AK
AMN
AN
Anatolia
Antiquity
ANYA
AP
APa
American Anthropologist, Washington
Archivio per TAntropologia e la Etnologia, Florence
Anthropologischer Anzeiger, Stuttgart
Archaeological Bulletin, New York
Acta Genetica et Statistica Medica, Basel
Annals of Eugenics, London
Abhandlungen zur exakten Biologie, Berlin
Journal of the International Institute of African Lan-
guages and Cultures, London
Acta Geneticae Medicae et Gemellologiae, Rome
Archives de I’Institut de Paleontologie Humaine, Paris
American Journal of Anatomy, Philadelphia
American Journal of Human Genetics, Baltimore
American Journal of Physical Anthropology, Philadel-
phia
American Journal of Science, New Haven
Animal Kingdom, New York
American Museum Novitates, New York
American Naturalist, Lancaster, Pa.
Revue annuelle de Tlnstitut d’Archaeologie de I’Uni-
versite d’ Ankara, Ankara
Cambridge, England (formerly Newbury, Berkshire)
Annals of the New York Academy of Sciences, New
York
Asian Perspectives, Hong Kong
Annales de paleontologie, Paris
xxxvi
APAM
AQ
AR
ARSI
AS
ASAG
ASAM
ASPR
ATM
AuS
RAM
BAMN
BASP
BBMNH
Belleten
BGI
BGSC
BIAF
Biometrika
BMFM
BMSA
BPGO
BRCI
BS
BSA
BSGI
BSPC
BSPF
BUM
Periodicals and Their Abbreviations
Anthropological Papers of the American Museum of
Natural History, New York
Anthropological Quarterly, Washington, D.C.
Anatomical Record, Philadelphia
Annual Report of the Smithsonian Institution, Wash-
ington, D.C.
Archives des sciences physiques et naturelles, Geneva
Archives suisses d’ Anthropologie g enerale, Geneva
Annals of the South African Museum, Cape Town
American School of Prehistoric Research, Cambridge,
Mass.
Annals of the Transvaal Museum, Pretoria
The Australian Scientist, Sydney
Bulletin d’Archeologie Marocaine, Rabat
Bulletin of the American Museum of Natural History,
New York
Bidletin of the American School of Prehistoric Re-
search, Cambridge, Mass, (formerly New Haven)
Bulletin of the British Museum of Natural History,
London
Belleten Turk Tariha Kururnu Basimevi, Ankara
Bollettino della Societa geologica italiana, Rome
Bidletin of the Geological Society of China, Peking
Bulletin d Plnstitut francais d’Afrique Noire, Paris
London
British Museum Fossil Mammals of Africa, London
Bulletins et Memoires de la Societe d’ Anthropologie,
Paris
Beitrage zur Paldontologie und Geologie Oesterreich-
Ungarns und des Orients, Vienna
Bulletin of the Research Council of Israel, Jerusalem
Bulletin Scientifique, Conseil des Academies de la RPF
de Yugoslavie, Zagreb.
Bulletin de la Societe d’ Anthropologie de Paris, Paris
Bulletin du Service Geologique de Vlndochine, Hanoi
Boletim da Sociedade P ortuguesa de Sciencias Natu-
rais, Coimbra
Bulletin de la Societe Prehistorique Franqaise, Paris
Bulletin of the University Museum, University of Penn-
sylvania, Philadelphia
CA
CH
CHM
Circulation
CIWP
CMES
CMJ
CNHS
CRAS
CSHS
Cytologia
DAKM
Diogenes
DR
Endocrinol-
ogy
ER
Evolution
Expedition
Experjentia
FICA
GHSP
GS
HAS
HB
IGC
IJNS
ILN
IVPM
Periodicals and Their Abbreviations xxxvii
Current Anthropology, Chicago
Collection Hesperis, Paris
Cahiers d’Histoire Mondiale, Paris
New York
Carnegie Institution of Washington Publications,
Washington, D.C.
Ceylon Museum Ethnographic Series, Colombo
Chinese Medical Journal, Shanghai
Ceylon National Museums Natural History Series, Co-
lombo
Comptes-rendus Hebdomadaires des Seances de T Aca-
demic des Sciences, Paris
Cold Spring Harbor Symposia on Quantitative Bi-
ology, Cold Spring Harbor, New York
Tokyo
Deutsches Archiv fur klinische Medizin, Leipzig
Chicago
The Dental Record, London
Los Angeles
Eugenics Review, London
Hempstead, New York
Philadelphia
Basel
Fifth International Congress of Anthropological and
Ethnological Science, 1956, Philadelphia
Geologica Hungarica, Series Paleontologica, Budapest
Gottinger Studien, Gottinger
Harvard African Studies, Cambridge, Mass.
Human Biology, Detroit
International Geological Congress, London, 1948
Indonesian Journal for Natural Science, Jakarta
Illustrated London News, London
Institute of Vertebrate Paleontology Memoirs, Peking
xxxviii
TAnat
JAP
JCPP
JEAN
JFS
JGen
JJP
JMBR
JPh
JPLS
JPSI
JRAI
JRAS
L’anth
Man
Mankind
MB
MJA
MKNA
MNGB
MNMM
MOG
MRSE
Nature
NC
NG
NH
P eriodicals and Their Abbreviations
Journal of Anatomy, London
Journal of Applied Physiology, Washington, D.C.
Journal of Comparative and Physiological Psychology,
Baltimore
Journal of the East Africa Natural History Society,
Nairobi, Kenya
Journal of Forensic Sciences, Chicago
Journal of Genetics, Cambridge, England
Japanese Journal of Physiology, Nagoya
Journal of the Malayan Branch of the Royal Asiatic
Society, Singapore
Journal of Physiology, London
Journal of the Proceedings of the Linnaean Society of
London, London
Journal of the Paleontological Society of India, Luck-
now
Journal of the Royal Anthropological Institute of Great
Britain and Ireland, London
Journal of the Royal Asiatic Society of Great Britain
and Ireland, Ceylon Branch, Colombo
L’Anthropologie, Paris
London
Sydney, Australia
Monographiae Biologicae, The Hague
Medical Journal of Australia, Sydney, Australia
Mededelingen Koninklijke Akademie van Wetenschap-
pen, Amsterdam
Mitteilungen der Naturforschenden Gesellschaft in
Bern, Bern
Memoires of the National Museum of Melbourne, Vic-
toria, Australia
Medelelser om Gronland, Copenhagen
Memorias Real Socieclad Espahola de Historia Natural,
Madrid
London
Neanderthal Centenary, Utrecht
National Geographic Magazine, Washington, D.C.
Natural History, New York
Periodicals and Their Abbreviations
XXXIX
NMRI Naval Medical Research Institute Lecture and Review
Series, Bethesda, Md.
NYT The New York Times
Oceania
OJS
PAf
PAPS
PASA
PBM
PKAW
PKSF
PLSL
PMP
PNAS
PNHB
PPS
PRSM
PS— D
PS— NS— D
PTCPFA
PTPA
PYMP
Sydney, Australia
Ohio Journal of Science, Columbus
Paleontologia Africana, Johannesburg
Proceedings of the American Philosophical Society,
Philadelphia
Proceedings of the Academy of Science of Amsterdam,
Amsterdam
Perspectives in Biology and Medicine, Chicago
Proceedings: Koninklijke nederlandse akademie van
Wetenschappen, Amsterdam
Publications under the Keith Sheridan Foundation for
Medical Research, Adelaide, S. Australia
Proceedings of the Linnaean Society of London
Peabody Museum Papers, Cambridge, Mass.
Proceedings of the National Academy of Science,
Washington, D.C.
Peking Natural History Bulletin, Peking
Proceedings of the Prehistoric Society, Cambridge
Proceedings of the Royal Society of Medicine, London
Paleontologia Sinica, Series D, Peking
Paleontologia Sinica, New Series D, Peking
Proceedings of the Third Congress on the Prehistory
of the Far East, 1938
Proceedings of the Third Pan-African Congress on
Prehistory held in N. Rhodesia, Livingstone, 1955,
pub. London, 1957
Postilla Yale Peabody Museum, New Haven, Conn.
QJGS Quarterly Journal of Geological Sciences, London
QRB Quarterly Review of Biology, Washington, D.C.
Quartar Bonn
Quaternary Rome
RA Rivista di Antropologia, Rome
RBMO Research Bulletin, University of Missouri, College of
Agriculture, Agricultural Research Station, Co-
lumbia, Mo.
xl
Periodicals and Their Abbreviations
RGA
RM
RQS
RR
RSAM
Revista Geografica Americana, Buenos Aires
Richerche di Morfologia, Rome
La Revue des Questions Scientifiques, Louvain
Radical Review, New Bedford, Mass.
Records of the South Australian Museum, Adelaide,
S. Australia
SA
SAAB
SAJS
Science
SD
SE
SIAr
SM
SMC
SMJ
SNNM
SRP
SSF-CB
Sumer
SVfZ
SWJA
SZC
Scientific American, New York
South African Archaeological Bulletin, Cape Town
South African Journal of Science, Johannesburg
Washington, D.C.
Science Digest, Chicago
Sovetskaia Etnografiia, Moscow
Slovenska archeologia, Brno
Saugetierkundliche Mitteilungen, Stuttgart
Smithsonian Miscellaneous Collections, Washington,
D.C.
Saraivak Museum Journal, Kuching, Borneo
Sodlogiese naborsing nasionale Museum, Bloemfontein,
S. Africa
Smithsonian Report, Publication, Washington, D.C.
Societas Scientiarum Fennica, Commentationes Biolo-
gicae, Helsinki
Baghdad
Schweizerische Viertelfahrsschrift fur Zahnheilkunde,
Zurich
Southwestern Journal of Anthropology, Albuquer-
que, N.M.
Spolia Zeylanica, Colombo
TB
The Leech
TI
TIBS
TLAB
TMM
TNYA
Triangle
Tabulae Biologicae, The Hague
Johannesburg, S. Africa
The Interamerican, Denton, Texas
Trabajos del Institute “Bernadino de Sahagun” de
Antropologia y Etnologia, Madrid
Travaux du Laboratoire d’Anthropologie et d’Archeo-
logie Prehistoriques du Musee du Bardo, Algiers
Transvaal Museum Memoires, Pretoria
Transactions of the New York Academy of Sciences,
New York
Basel
Periodicals and Their Abbreviations
xli
TRSL
TRSS
UMM
VGR
VGPA
VP
WADC
WMDM
YAPS
ZA
ZfE
ZfMuA
ZfNF
ZfRK
ZOOLOGICA
zv
ZZ
Transactions of the Royal Society of London, London
Transactions of the Royal Society of South Africa,
Cape Town
University Museum Memoirs, University of Pennsyl-
vania, Philadelphia
Verhandlungen der Geologischen Bundesanstalt, Vi-
enna
V erhandlungen der Gesellschaft fur physische Anthro-
pologie, Stuttgart
Vertebrata Paleasiatica, Peking
Wright Air Development Center Technical Reports,
Wright-Patterson Air Force Rase, Ohio
W etenschappelijke Mededelingen Dienst van de Mijn-
bouw, Bandoeng
Yearbook of the American Philosophical Society, Phila-
delphia
Zoologischer Anzeiger, Leipzig
Zeitschrift fiir Ethnologie, Berlin
Zeitschrift fiir Morphologie und Anthropologie, Stutt-
gart
Zeitschrift fiir Naturforschung, Tubingen
Zeitschrift fiir Rassenkunde, Stuttgart
New York
Zoologische Verhandelingen, Leyden
Zinruigaku Zassu, Journal of the Anthropological So-
ciety of Nippon, Tokyo University
THE ORIGIN OF RACES
a
1
THE PROBLEM
OF RACIAL ORIGINS
A On the Antiquity of Races
t the dawn of history, which is another way of saying
beginning with Herodotus,” literate people of the ancient world
were well aware that mankind was divided into a number of
clearly differentiated races. Even before that, racial differentia-
tion can be traced back to at least 3,000 b.c., as evidenced in
Egyptian records, particularly the artistic representations. We also
have pictures of white people on the walls of western European
caves which are as much as 20,000 years older.
How many kinds of people there were in the world was not
really known until after the voyages of discovery that tore the veil
from the Americas, the Pacific islands, and Australia. Even then
the problem of classifying the races remained, and it has not been
settled to this day.
For present purposes I am using a conservative and tentative
classification of the living peoples of the world into five basically
geographical groups: the Caucasoid, Mongoloid, Australoid, Con-
goid, and Capoid. The first includes Europeans and their overseas
kinsmen, the Middle Eastern Whites from Morocco to West Paki-
stan, and most of the peoples of India, as well as the Ainu of Ja-
pan. The second includes most of the East Asiatics, Indonesians,
Polynesians, Micronesians, American Indians, and Eskimo. In the
third category fall the Australian aborigines, Melanesians, Papu-
ans, some of the tribal folk of India, and the various Negritos of
South Asia and Oceania. The fourth comprises the Negroes and
4 The Problem of Racial Origins
Pygmies of Africa. I have named it Congoid after a region ( not a
specific nation) which contains both kinds of people. The term
Negroid has been deliberately omitted to avoid confusion. It has
been applied both to Africans and to spiral-haired peoples of
southern Asia and Oceania who are not genetically related to
each other, as far as we know.1 Negroid will be used in this book to
denote a condition, not a geographical subspecies. The fifth group
includes the Bushmen and Hottentots and other relict tribes, like
the Sandawe of Tanganyika. It is called Capoid after the Cape of
Good Hope. If this subspecies once occupied Morocco (see Chap-
ter 13), the cape can be thought of as Cape Spartel. Either way,
the term is appropriate.
My aim in this book is to see how far back in prehistoric an-
tiquity these human racial groups can be traced. Did they all
branch off a common stem recently, that is, within a few tens of
thousands of years, after mankind had evolved as a single unit to
the evolutionary state of the most primitive living peoples? Or did
their moment of separation lie lower down on the time scale, when
long-extinct types like the so-called ape men of Java and China
were still alive? If the second is true, much of the evolution of the
different existing races may have taken place separately and in
parallel fashion over a period of hundreds, rather than tens, of
thousands of years. The first hypothesis is the one more commonly
held, but it presents some impressive stumbling blocks.2
If all races had a recent common origin, how does it happen
that some peoples, like the Tasmanians and many of the Australian
aborigines, were still living during the nineteenth century in a
manner comparable to that of Europeans of over 100,000 years
ago? Either the common ancestors of the Tasmanians cum Aus-
tralians and of the Europeans parted company in remote Pleisto-
cene antiquity, or else the Australians and Tasmanians have done
some rapid cultural backsliding, which archaeological evidence
disproves.
If the ancestors of the living races of mankind were a single
1 They differ completely in blood-group patterns, particularly in the Rhesus
genes.
2 See W. W. Howells, Jr.: Mankind in the Making (New York: Doubleday and
Company; 1959), especially p. 236; and C. S. Coon’s review of same in Science,
Vol. 130, No. 3386 (1959), pp. 1399-1400.
On the Antiquity of Races
5
people a few thousands of years ago and they all spoke a single
language, how does it happen that the world contains thousands
of languages, hundreds of which are unrelated to each other, and
some of which even use such odd sounds as clicks? Some lan-
guages are tonal and others are not, and the difference between a
tonal and a nontonal language is basic and profound. Eskimo and
Aleut, which are closely related languages, have been separated
for about two thousand years. It takes at least twenty thousand
years for two sister languages to lose all semblance of relation-
ship.3 If, therefore, all languages are derived from a single
mother tongue, the original separation must go back many times
that figure. The only alternative is that more than one line of
ancestral man discovered speech independently. Even so, the
number of languages spoken by a single subspecies, the Mongo-
loid, is great enough to imply a vast antiquity.
All the evidence available from comparative ethnology, linguis-
tics, and prehistoric archaeology indicates a long separation of
the principal races of man. This is contrary to the current idea
that Homo sapiens arose in Europe or western Asia about 35,000
b.c., fully formed as from the brow of Zeus, and spread over the
world at that time, while the archaic species of men who had
preceded him became conveniently extinct. Actually, the homines
sapientes in question were morphologically the same as living
Europeans. To derive an Australian aborigine or a Congo Pygmy
from European ancestors of modern type would be biologically
impossible.
The current idea is based on the study of comparative anatomy
without reference to evolution, and a misunderstanding of pale-
ontology. One anatomist, Morant,4 found by means of a number of
measurements taken on less than ten Neanderthal skulls that this
ancient population differed in mean measurements from a number
of modern populations more than the modern skulls differ from
each other. The differences reflected mainly the fact that Nean-
3 D. H. Hymes: “Lexicostatistics So Far,” CA, Vol. 1, No. l (i960), p. 3-44.
The 20,000-year calculation, a conservative figure, is my own, based on Hymes’s
data.
4 G. M. Morant: “Studies of Paleolithic Man, II, A Biometric Study of Nean-
derthaloid Skulls and of Their Relationships to Modern Racial Types,” Biometrika,
Vol. 2 (1927), pp. 318-80.
° The Problem of Racial Origins
deithal men had low, flattish cranial vaults and protruding faces;
but these features could have come from a small number of genes
concerned with adaptation to cold weather. Since 1927, when
Morant’s study was published, “progressive” and “transitional”
high-headed Neanderthals have been unearthed in western Asia.
These new discoveries suggest that the total extinction of that
fossil race is unlikely. We now have fossil skulls from China, Af-
rica, and Europe, found since Morant studied the Neanderthals,
which closely resemble the modern races in features that seem to
have evolved and been handed down locally. Such features in-
clude the extent to which the face is flat or beak-like, the shape of
the nasal bones, and the size ratio of front teeth to molars. If we
grant that races, like the species to which they belong, can evolve,
our problem becomes simpler.
The misinterpretation of paleontology by nonpaleontologists
came about naturally. Anyone who studies the family trees of
various lines of animals over millions of years is bound to be im-
pressed by the multitude of extinct species, and to notice that the
living animal species are descended from very few ancestral ones.
When this observation is applied to many forms of life over the
span of geological time, it holds true; but for man it does not. Man
is little more than a half million years old. Geologically speaking,
we were born yesterday. The fossil men now extinct differed
from each other in race, and were not members of separate species
except in the sense that one species grew out of another.
As human beings are animals, they are subject to the same laws
of evolutionary change that govern the rises and falls of other
species and their transmutations into increasingly complex and
efficient forms. Therefore we have two jobs to do: (1) to survey
the rules of species formation and the differentiation of races, in-
cluding the composition of populations, systems of mating, dif-
ferential fertility, and geographical adaptation at different ecolog-
ical levels, as they may apply to man; and (2) to go over with a
fine-toothed comb all the original evidence about fossil specimens
of man and his predecessors which can be found. This includes
actual specimens, casts, and technical reports, some lying on the
bottom shelves of library stacks, with pages still uncut, and un-
disturbed for decades. Because few textbook writers have both-
The Problems of Human Taxonomy: the Genus g
ered to consult these primary sources, few new ideas about the
evolution of races have reached the public for a long time.
The Problems of Human Taxonomy: the Genus
Over two hundred years ago Linnaeus, the father of taxonomy,5
or systematics as he called it, initiated the practice of giving each
species in nature an italicized double name, or binominal, one of
which was Homo sapiens. The first word is the name of the genus
and the second that of the species itself. In the species Homo
sapiens he included all living peoples. At that time no fossil men
had been discovered, and the genus Homo had therefore but a
single species.
Linnaeus used only one word to designate biological units
smaller than the species: variety. At that time the concept had
not yet arisen that the unit of inheritance and evolution is the
population to which an individual belongs rather than the indi-
vidual himself, and the exact meaning of variety was not clear. In
recent years taxonomists, in reviewing the nomenclature of spe-
cies, have found that many units given specific rank in the past
were subspecies, or geographical races, of larger units, and that
what had been called varieties were races of one magnitude or
another, or even individual variants.
In order to obtain material for classification, zoologists were
kept busy collecting skins and skulls of many kinds of animals,
and paleontologists removing bones, teeth, claws, and shells of
ancient animals from the ground. Rarely did the paleontologists
have whole skeletons to work with; and even when they did,
characteristics studied by zoologists, such as hair form and color,
skin structure, and the number of mammary glands, could not be
determined except in a very few cases, as when mammoths were
found frozen in the ground.
Whereas zoologists could collect large numbers of contempo-
rary specimens, paleontologists sometimes possessed only unique
For a lucid introduction to this subject, see G. G. Simpson: Principles of
Animal Taxonomy (New York: Columbia University Press; 1961) and “The
Principles of Classification and a Classification of Mammals,” BAMN, Vol. 85
(i945)-
10 The Problem of Racial Origins
specimens, which had to be related to others from different times
and different places. Often the time gap between apparently re-
lated specimens was so great that it was unlikely that they could
have belonged to a single species. Being cautious men, most
paleontologists considered it more conservative to give separate
generic names to unique or rare fossils of different periods than
to assume their identity, particularly when in living animals such
as the sheep and goat, which belong to different genera, the only
difference visible in the skelton is the relative lengths of the seg-
ments of the forelimb. Paleontologists therefore formed the habit
of giving new and unique specimens separate generic names,
setting aside the finer classification of related species until more
bones had been found.
When, in the second half of the nineteenth century, paleontolo-
gists and archaeologists began turning up the bones of fossil men,
some of them applied this practice to the much more limited field
of anthropology, and we find such designations as Pithecanthro-
pus erectus, Sinanthropus pekinensis, and more recently, Atlan-
thropus mauretanicus tagged to specimens some of which differ
from one another no more than do individuals in the living species.
Homo sapiens
The final difficulty with this type of taxonomy is that it can-
not be reconciled with our time scale. Simpson, Kurten, and others
have shown that, within the geological periods with which we are
concerned, a genus of mammals requires about eight million years
to establish itself, and it usually makes no difference whether the
animals are large or small, or fast or slow to mature.6
The oldest fossil-man remains that are definitely and indu-
bitably Homo may be no more than 700,000 years old. If there
really were, during the last 700,000 years, four genera of fossil
men, including Homo, Pithecanthropus, Sinanthropus, and Atlan-
6 Simpson: The Major Features of Evolution (New York: Columbia University
Press; 1953).
B. Kurten: “Rates of Evolution in Fossil Mammals,” CSHS, Vol 24 (1959)
pp. 205-15.
11
The Species Concept
tkropus, then these genera must have parted company early
in the Pliocene, and we have neither manlike bones nor tools from
this period.
Later on, after tools had appeared, we find that both Atlan-
thropus in North Africa and Homo in Europe were making stylis-
tically similar stone implements. Although a great many claims
can be made for parallel evolution, it is inconceivable that men of
two distinct genera could have made similar tools.
The concept that the fossil men so far found, who lived during
the last half million years, belonged to more than one genus is
impossible both anatomically and in terms of behavior, as re-
vealed by archaeology. This concept must be abandoned, and
indeed many zoologists and anthropologists have already dis-
carded it. Of the names proposed for our genus, Homo has two
centuries of priority, and Homo is what we are, what our known
ancestors were, and what our unknown ancestors could have been
for as long as eight million years.
The Species Concept
In the whole field of taxonomy no identification is as impor-
tant as that of the species of an animal. Higher categories, such as
the genus, family, order, and so on, are subject to argument and
revision, and lower categories, the subspecies and local race, are
also more difficult to establish. The species, however, is the pivot
of the entire structure because it is the unit of evolutionary
change.
In the early days of taxonomy, a collector would shoot a bird
or animal, keep its skin and skull, compare it with others in exist-
ing collections to determine whether it was something new, and
if it was, he would write up a detailed description, giving the
bird or animal a new name. It thus became the type specimen, or
holotijpe, of its species, and future collectors would compare their
discoveries with it. This practice was applied to the anthropologi-
cal field. Blumenbach, whose classification of mankind in the
familiar fivefold skin-color system is still used in some school geo-
1 2 The Problem of Racial Origins
graphy books, selected a particularly handsome skull from a Eu-
ropean collection as the type specimen of the white race, and as it
had belonged in life to a native of the Caucasus Mountains, white
people came to be called Caucasians, or Caucasoids, and still are.
As late as 1912 Boule selected the skeleton of La Chapelle aux
Saints as the type specimen of Neanderthal man, which he com-
pared to the skeletons of one Frenchman and three anthropoid
apes.
As early as Darwin, however, it was recognized that a species is
not just the specimen that happened to be killed or unearthed
first, and others later found to resemble it, but a population. In-
deed, Darwin based his theory of natural selection on his obser-
vation that individuals of a species are variable, and that one
need not be more typical than another. As time went on, it became
clear that a species is a breeding unit or population, which has a
gene pool of its own, and not just a collection of individuals, and
that each population is a separate entity, living in two related
states of dynamic equilibrium. The first regulates the balance be-
tween the individuals that compose the population. The second
governs its relations with the other species in its environment.
Another early observation was that members of different spe-
cies do not interbreed, at least in a state of nature. It was first
thought that this was not for lack of trying but simply because
each species was incapable of fertility with any other. However,
early in the twentieth century the rising science of genetics made
it clear that some animals of different species could produce fer-
tile offspring if they could be made to come together. Sterile hy-
brids like the mule were known from antiquity, and tiger-lion mix-
tures have been produced in zoos, but hybridization, it was found,
is not a common or important mechanism of evolutionary change
in the higher animals, as it is in plants. Furthermore, as each spe-
cies is in genetic equilibrium with its environment, the addition of
new genes from an animal with a different kind of equilibrium
could be expected to produce offspring less viable than either
parent.
The important distinction is that members of potentially inter-
fertile species do not ordinarily interbreed either because their
13
The Species Concept
breeding periods fall at different seasons or because they simply
do not attract each other: they do not recognize each other’s mat-
ing symbols — visual, olfactory, auditory, or whatever.
In any case, whether or not unconfined animals of different
populations interbreed when given the opportunity is the critical
test of a zoological species. Paleontologists, of course, cannot use
this test, which may be another reason why they prefer to deal in
the more readily identified unit of the genus. In the case of living
human populations, we can confirm Linnaeus’s decision that all
men belong to the same species, not only because all races are in-
terfertile but also because some individuals among them inter-
breed, although others oppose mixture. In the case of early human
populations unearthed by archaeologists, we cannot be sure
whether interbreeding has or has not taken place; and at only one
site, the Mt. Carmel caves of Palestine, is there any evidence — a
high degree of individual variability combined with a mingling of
tool forms — to suggest that the races were mixing, but even that is
inconclusive. Therefore, the statements commonly made that
Pithecanthropus, Sinanthropus, Neanderthal man, or a member
of any other ancient population was unable to interbreed with his
neighbors, if he had any, is speculative and cannot be demon-
strated.
These statements are based on the old idea that if in some char-
acteristic the ranges of variability of two populations fail to over-
lap, then these populations are different species. If this were true,
then the Pygmies and Watusi of Ruanda-Urundi in Central Af-
rica, who live near each other, would be different species on the
basis of stature, and the black-skinned and white-skinned races of
the world would also be different species.
This obsolete concept of single-character taxonomy has long
since been abandoned. Zoologists now base their decisions on all
the characteristics they can identify and measure, characteristics
which together give the animal its essential nature, its (to borrow
a psychological term) gestalt. The determination of species can-
not be made by feeding figures into a computer. It is in a sense an
art, practiced by men of experience who know, first of all, how
species are formed.
14
The Problem of Racial Origins
The Spatial Requirements of Species
and Their Geographical Differentiation
Zoologists recognize two kinds of species, monotypic and
polytypic.7 A monotypic species contains a single pattern of ge-
netic composition, usually because it is a single population that
occupies a single, environmentally unified lebensraum in which
interbreeding is easy from one end of its territory to the other.
Monotypic species are in the minority. A polytypic species, on the
other hand, is broken up into a number of separate populations,
each occupying its own territory. Usually these territories adjoin
each other but are partially separated by environmental barriers.
Gene flow across the barriers is infrequent enough to permit the
development of separate genetic patterns but frequent enough to
prevent the different populations from becoming individual spe-
cies. When these barriers become absolute, local speciation can
occur. Once a new species has arisen, it is likely to expand into a
number of territories, where adaptation to new conditions will be
rapid. This is undoubtedly what happened to our ancestors once
they had acquired the erect posture and begun to use their hands
for something beside locomotion and their mouths for something
other than feeding and biting.
Regional populations of a polytypic species, once it has become
established and has spread, are normally allopatric, a term which
means simply occupying different territories.” If they were not
allopatric, they would compete with each other for food, and one
would drive out or absorb the other. Normally the one longest in
situ has the advantage over newcomers because it has adapted it-
self to its new environment by favorable genetic changes, unless a
geographical principle is involved, as in the case of isolated popu-
lations like those that arise on islands. Because they evolved with-
out competition, such populations are usually vulnerable when
This term should not be confused with the word polymorphic , which means,
in the language of geneticists, that inside a given population more than one gene
is available for a given position on a chromosome; the father may carry one, & the
mother another. The best-known example is the possibility of having a gene for
A, B, or O on a single chromosome in the ABO blood-group system.
The Subspecies
15
their territories are invaded by newcomers which evolved on large
continental areas where competition is keen.
Related species, however, can be sympatric, which is zoologese
for saying that they can occupy a single territory without inter-
fering with each other, just as zebras, wildebeeste, and giraffes
feed together on an African plain. Sympatric occupation is the
rule for animals that belong to different genera, families, orders,
and even higher categories of classification, which is why we have
regional faunas. It is not very common among closely related spe-
cies because they usually compete for food.
Whether or not related species are sympatric or allopatric de-
pends to a large extent on their eating habits. If a species special-
izes in a narrow dietary range, it can coexist with another that
specializes in a different range. The Australian koala lives essen-
tially on the leaves of a few kinds of eucalyptus, the presence of
which limits its range but allows it to coexist with other species of
marsupials on the ground below; the giant panda of western
China subsists largely on bamboo shoots whereas the smaller red
panda eats a variety of foods.
Animal species that specialize in food are called stenophagous,
the Greek term for narrow-feeding. Those that eat many kinds of
food are called euryphagous, or wide-feeders. Like any other spe-
cialty, stenophagy permits a rapid expansion in a narrow milieu,
but it is not the road to evolutionary success. Euryphagy involves
an animal in heavy competition, but if it survives, it has a better
chance of expanding over areas with differing food supplies, and
of undergoing further speciation.
In the case of man, he is euryphagous and always has been.
Man can eat roots, succulent leaves, fruits, berries, eggs, and flesh.
Except for grass, he can eat virtually everything that other ani-
mals eat, and this puts him in competition with many other spe-
cies and with other populations of his own and related species.
The Subspecies
The next taxonomic division below that of species is the sub-
species. A subspecies is a regional population of a polytypic spe-
*6 The Problem of Racial Origins
cies (a species with a number of separate populations) which
meets two tests: (1) it occupies a distinct geographical territory ;
(2) it differs from other subspecies of the same species in measur-
able characteristics to a considerable degree (to be specified
shortly).
Subspecies must by definition be allopatric: if several subspe-
cies were to inhabit a single region, they would breed together
and the differences between them would be obliterated. Within
its own geographical territory, which has an environmental char-
acter of its own, the subspecies has achieved, or is in the process
of achieving, an adjustment to its local food supply, to the local
climate, and to the behavior patterns of other animal species with
which it shares its domain. After each subspecies has worked out
a balance with all other elements in its local environment, it is not
likely to change very much until its situation changes: natural se-
lection will prune off unfavorable mutations that arise locally and
keep the favored gene ratio constant.
Over the border, which may be a natural barrier such as a range
of mountains or a patch of desert, or even a critical isotherm, may
be found another subspecies of the same species, equally well es-
tablished in a state of equilibrium with its environment. As the
two environments differ in certain details, so do the genetic struc-
tures of its occupants. What is good for A is less advantageous for
B, and vice versa. In each territory, natural selection keeps the
gene structure of the local subspecies constant by also eliminat-
ing unfavorable genes that flow over the border. However, genes
which are unfavorable in both environments may be eliminated
in both populations, so that A and B may evolve together into a
new polytypic species that retains its original set of subspecies.
This is what we think happened when a number of human sub-
species passed the threshold from Homo erectus to Homo sapiens.
Taxonomists have set up an arbitrary procedure to determine
whether two or more populations within a species are morpho-
logically different enough to qualify as subspecies. It is called the
overlap test and is applied both to visible criteria, such as tooth
size, and to invisible ones, such as blood groups. If in any well-
defined, presumably heritable morphological character, a repre-
sentative sample of population A differs from a representative
17
Mosaics, Clines, Local Races, and Racial Types
sample of population B to or beyond a critical degree, then we are
dealing with subspecies. The critical degree is 75 per cent. If 75
per cent or more individuals of A are different from 100 per cent of
B, then the two are probably subspecies.8
This method was devised for use on large samples of living ani-
mal populations and it can be applied to modern anthropometric
series, but it is rarely if ever useful in the study of fossil man be-
cause we have few samples large enough for analysis by proba-
bility statistics. When applied to modern human populations, this
test shows that Homo sapiens is at present a polymorphic species
divided into a number of clearly differentiated subspecies, each
centered in its own territory.
The concept of subspecies is essentially zoological and is used
almost entirely to describe regional variations in animal species.
However, paleontologists also use it occasionally, to describe steps
in a single evolutionary line which they consider too small to
merit the rank of separate species. Such units may be called suc-
cessional subspecies, or waagenons — named for a mid-nineteenth-
century paleontologist, W. Waagen.9 In order to keep confusion to
a minimum I shall not use the word subspecies in this book to
designate such successive units. When successive species must be
split, I shall do it in terms of the evolutionary levels or grades
through which they have passed.
Mosaics, Clines, Local Races, and Racial Types
Below the taxonomic level of the subspecies, zoologists find a
sometimes bewildering array of local racial variations of a minor
nature, which exist because subspecies as well as species can be
polytypic. This is as true of men as it is of mice, for man is the
8E. Mayr, E. G. Linslev, and R. L. Usinger: Methods and Principles of Sys-
tematic Zoology (New York: McGraw-Hill Book Company; 1953), p. 146. When
biometric statistical constants are available, this test can be performed without
plotting frequency curves by using the formula C.D. = ^ in which C.D.
equals Coefficient of Difference, Mi and M» the means of two series, and <r, and <r._.
their standard deviations. If the C.D. is 1.28 or higher, subspecific rank is indi-
cated. Attributes expressed in percentile values rather than means may be com-
pared directly.
9 Simpson: Principles of Animal Taxonomy, pp. 175-6.
The Problem of Racial Origins
most mobile of mammals. He walks the land, flies the skies, and
rides the oceans.
Part of the racial complexity of Homo sapiens disappears if we
disregard for the moment the distribution of modern peoples like
white and Negroid Americans, Latin Americans, South Africans,
and white Australians and New Zealanders, whose ancestors
reached their homes by ocean-going ships in recent times. Before
then each of the five subspecies recognized in this book was firmly
and uniquely installed in its geographical center. Between the nu-
clei of these five centers lie intermediate regions of two kinds.
One of them is the mosaic, which contains relict populations
living as enclaves in refuge areas. For example, in India at least
two forms of Australoids, classified as “tribal peoples,” dwell in
the hills, surrounded by Caucasoids whose home is the plains.
Such a mosaic pattern is the product of earlier, but not geologi-
cally ancient, migrations that have not had time to fuse. As will be
shown in the next chapter, it is typical of the tropics of the Old
World.
The other is a region of racial transition, a frontier-in-depth
within which a subspecies grades into another through intermedi-
ate forms. It may be called a clinal zone because in it the popula-
tion of the species intergrades in one or more measurable charac-
ters. In each heritable feature, the gradient is called a cline.1 For
example, the living Europeans grade from a high frequency of
blue eyes in the northwest, particularly in Ireland and Scandina-
via, to a high frequency of brown eyes in the southeastern part of
the continent. This eye-color gradient is a cline.
Whole complexes of related clines are found in clinal zones.
For example, in central Asia north of the Himalayas Caucasoids
merge into Mongoloids through the persons of several Turkic-
speaking peoples like the Kirghiz, Uzbeks, and Turkomans. This
clinal zone is a broad one. On the southern face of the Himalayan
wall a similar but narrow clinal zone stretches through a steep in-
termediate altitude zone, in northern India, Nepal, Sikkim, Bhu-
tan, and NEFA (Northeast Frontier Agency). As can be seen by
1J' S- Huxley: “Clines: An Auxiliary Taxonomic Principle,” 'Nature, Vol. 142
(1938), p. 219. See also Simpson: Principles of Animal Taxonomy, pp. 178-80.
Mosaics, Clines, Local Races, and Racial Types lg
these examples, the sharper the environmental barrier the nar-
rower the clinal zone between subspecies.
Not only in relict enclaves and clinal zones, but also within the
nuclear territories of subspecies, regional populations of minor
rank may be found which differ from each other in perceptible
ways short of the requirements of subspecies. These are known as
local races. As they rise and disappear rapidly, they receive little
attention from zoologists and usually none from paleontologists.
In man they are considered important by people without a bio-
logical background, usually because such groups may be identi-
fied to a certain extent with social, political, or religious units.
How many local races could be identified and counted among
living men is difficult to say, and different anthropologists might
each find a different number. Such details are of no importance in
this book, but it is important for us to know that local races exist
and are formed by the same biological mechanisms that have
fostered larger taxonomic units in the past.
Races like the Nordic, Alpine, Mediterranean, East Baltic, and
Dinaric, which loom large in the Europe-centered literature of
anthropology, are neither subspecies nor, in a strict sense, local
races, although some local races may be defined in these terms.
These words have also been used in the sense of tijpes, which can
be picked out of local populations. One may find a Spaniard who
is typically Nordic in the midst of a population of Mediterra-
neans, including his own brothers. In a sense the situation is
genetically comparable to finding a man of blood group B whose
father’s group was A. Types selected in this fashion are interesting
to observe, and we notice them every day. Whether or not they
reflect the origins of a population in one way or another, we must
remember that from the taxonomic point of view such types are
not races but simply the visible expressions of the genetic varia-
bility of the intermarrying groups to which they belong.
However, if we return to the first test of subspecies, geographi-
cal integrity, we are at first sight on shakier ground. Whites, Ne-
groes, and American Indians occupy the United States sympat-
rically. Hindus, Fijians, and Europeans similarly occupy the Fiji
Islands, and many other examples might be cited. As we study
20
The Problem of Racial Origins
each instance, we find that this situation is a recent one, as time is
measured biologically, and it is always associated with the expan-
sion of peoples who have left the food-gathering stage of sub-
sistence far behind.
Let us omit, for the moment, the agricultural peoples of the
world and the colonists, and consider only the peoples who still
aie, or until recently were, food gatherers. These hunters and col-
lectors are drawn from all five geographical races listed on page 3.
Each race is confined to a single territory without overlap except
in two regions: India, and southeast Asia plus Indonesia. Owing
to a lack of skeletal material, we do not know when the ancestors
of the various food gatherers moved into India, nor indeed which
race was eailiest there. In southeast Asia and Indonesia we know,
as will be explained in Chapter 10, that Mongoloids began re-
placing Australoids about 10,000 years ago, after the invention of
the bow and the domestication of the dog had made some hunters
more efficient than others.
This southward movement was a trickle compared to what hap-
pened in many other places 4,000 years later. By or after 6,000 b.c.
a number of local populations began to advance from the ecologi-
cal niche of hunters and gatherers to that of food producers, and
territorial expansions followed. These movements started no more
than four hundred generations ago, counting twenty-five years to
a generation. The colonial movements that brought Europeans to
America, South Africa, Australia, and New Zealand took place
less than twenty-five generations ago; only about twelve genera-
tions separate most descendants of passengers on the Mayflower
from their celebrated forebears.
These various movements have greatly restricted the territo-
ries of aboriginal food gatherers, but gatherers are still present in
reduced numbers. Many more have been absorbed into the new
food-producing populations or have borrowed the techniques of
food production from newcomers to their territories. Since the be-
ginning of agriculture no new subspecies have arisen; the princi-
pal changes that have taken place have been vast increases in the
numbers of some populations and decreases to the threshold of ex-
tinction in others. All this points to one conclusion: the living sub-
species of man are ancient. The origins of races of subspecific rank
21
The Differentiation of Species
go back into geological antiquity, and at least one of them is as
old, by definition, as our species.
The Differentiation of Species
Species formation is believed to be the product of four
principal factors: mutation, recombination, selection, and isola-
tion.2 A mutation is a heritable, spontaneous, and within certain
limits random change in the chemical composition of a molecular
segment of a chromosome known as a gene or gene locus.3 These
changes take place normally in all organisms at individual fre-
quency rates that can be predicted. As most mutations produce
unfavorable effects, relatively few are passed on or participate in
species formation. The same mutation, favorable or otherwise, can
appear time after time, at its own rate, in individuals of different
races. Yet mutation is the primary element in evolution. The other
three are secondary.
Recombination, known as Mendel’s second law, is the process
by which rows of gene-molecules strung together on chromosomes
break up and form new associations.4 At meiosis, that critical mo-
ment in fertilization when a single array of paternal chromosomes
lines up with and joins a single set of maternal chromosomes, the
pairs do not always merge with each other in a regular fashion.
Some chromosomes cross over each other at various loci and trade
strings of genes. Others break up and the fragments attach them-
selves to other chromosomes or get lost. These new arrangements
can also cause changes in the resultant organism.
Selection is the well-known pruning process by which the envi-
ronment determines which novelty produced by mutation or re-
2 E. Mayr: “Change of Genetic Environment and Evolution,” in J. Huxley,
A. C. Hardy, and E. B. Ford, eds.: Evolution as a Process (London: George Al-
len and Unwin; 1954), pp. 157-80.
3 More technically, it is a change in the sequence of nucleotides within a DNA
( dexoribonucleic acid) molecule of a single chromosome. See P. Alexander:
“Radiation-Imitating Chemicals,” SA, Vol. 202, No. 1 (i960), pp. 99-108. Ac-
cording to Demarec, there are about 10 to 15 genes to each DNA molecule, or
20 to 30 to a pair of molecules. M. Demarec: “The Nature of the Gene,” AJHG,
Vol. 13, No. 1 (1961), pp. 122-7.
4 For present purposes this process is not also called a mutation. See Simpson:
The Major Features of Evolution, pp. 82-3.
22 The Problem of Racial Origins
combination shall gradually spread through the group because
of its superiority to the old trait it replaces, and which novelty
shall be eliminated because it is unfavorable. As most mutations
are unfavorable, when a species is not perceptibly changing, se-
lection serves almost entirely to preserve the status quo. However,
the process of replacement is characteristically slow. Old genes
have a habit of hanging on as minorities, and if the environment
changes back once more, they may re-emerge as majorities, in new
combinations.
Isolation, the fourth factor, is necessary for the rise of new spe-
cies because, unless a breeding population is self-contained, natu-
ral selection may be unable to eliminate old, unfavorable genes
from its pool. A constant gene flow from neighboring populations
may renew the old genes as fast as they are being lost. In a mono-
typic species such gene flow is impossible by definition. But in a
polytypic species only those genes can be eliminated which are
unfavorable to all its component units. When this happens, the
species evolves as a whole, whereas its component populations
may retain their local differences.
Balanced Polymorphism
Sometimes it is disadvantageous for a population to elimi-
nate its old genes completely. An old gene may possess the ability
to meet an old crisis, if that crisis should return. Furthermore, the
old gene and the new one with which it shares, as an alternate, its
position on a chromosome may do things together that neither
could do alone.
In genetic shorthand, AB may be better under some conditions
than either AA or BB. The best-known example of this effect in
man is probably the so-called sickling trait common among West
African Negroes. This is expressed by the letters S and s. S means
that you have the trait, s that you don’t. The S gene curls the red
corpuscles in the blood, impeding oxygen flow; the s gene has no
known effect. The S gene alone resists malignant malaria, which
kills many children. But an SS child may die of oxygen starvation,
and an ss child of malaria, whereas an Ss child is likely to survive
23
On the Timing of the Individual Growth Cycle
both diseases. The population profits by the retention of both
genes, each of which has a disadvantage in that particular envi-
ronment.
The example just cited may explain the presence of genetic
variability in many populations even though we don’t yet under-
stand why it is there in each case. It may also in part explain the
re-emergence of “types.”
On the Timing of the Individual Growth Cycle
In addition to mutation, recombination, selection, and iso-
lation, biologists have discovered a fifth evolutionary process
which is tertiary because it depends on combinations of the other
four, only one of which, mutation, is primary. This is a heritable
change in the time of appearance of different characters in the
growth cycle of the individual.
Each organism passes through three principal stages of devel-
opment. It starts as an embryo, a fertilized egg in the process of
cell division which has not yet reached the point where an em-
bryologist can tell its species. In man this condition lasts about
nine weeks. Then in mammals it becomes a fetus, in birds a chick,
and in insects a larva.5 After it has been born, pecked its way out
of its shell, or left its cocoon, it starts on the road to adult life in
different stages of preparation, depending on the class of animal it
belongs to.
Both in fetal and postnatal life, the individual must be adjusted
to its environment, or it will perish. Certain traits that are neces-
sary to the fetus and useless to an adult appear in fetal life and
then disappear. Other traits appear as they are needed. Inciden-
tally, it is not true that every individual recapitulates the forms of
all its ancestors from the beginning of life on earth. We do, how-
ever, recapitulate many of the fetal traits of our ancestors, but not
all of them, and not all in the original order. Nevertheless, the
fetus possesses a vast store of transient genetic characteristics that
could be used in adult life under different circumstances.
One of the features that all animals inherit is a built-in timing
5 G. R. de Beer: Embryos and Ancestors, 2nd. ed., (New York: Oxford Uni-
versity Press; 1951).
24
The Problem of Racial Origins
schedule which regulates the order of appearance and the dura-
tion of growth of different bodily systems. This schedule can be
upset through standard genetic mechanisms, such as mutation
and recombination. The survival of fetal traits into adult life occa-
sioned by such a change is called neoteny.
The classic example of neoteny is the life cycle of an amphibian
of the salamander group, the axolotl. This animal arrives at sexual
maturity during its tadpole stage and never leaves the water to
become an air breather like other salamanders, frogs, and toads,
but reproduces and dies in its original medium. Other examples
are found among certain birds that have lost the power of flight.
They retain throughout life the down that covers the chick before
it breaks out of its shell. Ostriches, emus, cassowaries, and pen-
guins have all acquired this neotenous change independently.
In man’s ancestors neoteny may have been at play before the
appearance of Homo erectus. The position of the head on the neck
at right angles to the axis of the vertebral column is neotenous; it
is found in the fetuses of all the primates and indeed in those of
other mammals. In the fetuses of primates in general the thumb is
relatively long in proportion to the length of the other fingers.
Among many monkeys and all apes the adult animals have short
thumbs, which in man remain neotenously long throughout life.
In insects, which are born fully grown and completely adult,
all changes in timing have to be neotenous. In mammals, which
are small when bom and dependent on their mothers for food and
protection, the infantile form differs markedly from the adult in
many ways. A baby mammal has to grow mightily and in most
species rapidly, and in the higher species it has much to learn. As
growth is largely controlled by the endocrines, any shift in endo-
crine balance can cause radical changes in the form and appear-
ance of the adult animal.
In man some laces appear infantile in certain respects
throughout life, whereas the children of other races look like
miniature adults. In some races the color of the hair never changes
during an individual s lifetime, except among persons who reach
advanced senility. In others the hair may start out blond, become
brown at puberty, and turn white by the age of thirty.
The classbook issued to the members of the Harvard class of
25
On Size and Form: Allometry
1925 at our twenty-fifth anniversary contains two portraits of each
man who was still alive in 1950 and who could be reached. One
portrait was taken at graduation, the other twenty-five years later.
In some individuals almost no change can be detected; others had
changed so much that they were unrecognizable. Yet nearly all
these men were of the same racial origin. Age changes, then, vary
within populations as well as between them. Not one of my class-
mates, however, looked like a Pygmy or a Bushman.
Races that retain a number of infantile features throughout life
are called pedomorphic; those in which mature features appear
early are called g erontomorphic, after the Greek words pais, a
child, and geron, an old man. Pedomorphism and gerontomorph-
ism are most conspicuous in external, visible anatomy, but they
can also affect the nervous system, the vocal cords, other covert
systems and structures, and behavior. Most fossil men that we
know were gerontomorphic, as witness their heavy brow ridges
and long faces. Homo sapiens as a whole seems to be relatively
pedomorphic, although variable in this respect both racially and
individually.
On Size and Form: Allometry
We must be careful, in seeking for relationships between dif-
ferent races, not to confuse pedomorphy and gerontomorphy with
normal variations that take place when animals of the same or re-
lated species grow smaller or larger. A mouse has a larger brain, in
proportion to its body size, than a rat does. A Great Dane’s eye-
balls are proportionately smaller, although absolutely larger, than
those of a terrier.
Animals that are otherwise genetically similar vary in propor-
tions according to size, the small ones being more compact, the
larger ones more attenuated. The principle governing these dif-
ferences is called allometry. Zoologists not only recognize this rule
but express it in formulas. For example, in the horse family face
length equals .3 times skull length, to the 1.2 power.13 A big horse
6 de Beer: op. cit., p. 27.
A classic work on this subject is D’A. W. Thompson’s Growth and Form (New
York: The Macmillan Company; 1945).
26
The Problem of Racial Origins
has a longer face, both absolutely and relatively, in proportion to
his skull length, than a small horse does. By the same token, an
average African Pygmy has relatively shorter legs and a relatively
larger head than does an average African Negro.
On Sexual Dimorphism
Another factor to be considered in comparing races and spe-
cies is the degree of differentiation between adult males and fe-
males in a population. This is called sexual dimorphism. It varies
greatly both in mammals and birds. Male and female cardinals
have feathers of different colors; yet it is difficult for a nonorni-
thologist to tell a male from a female robin. Among the primates,
a male gorilla may be twice as large as any member of his harem,
whereas the only visible difference in gibbons in the wild is the
protrusion, through the fur, of nipples in the female that has
borne offspring.
Sexual dimorphism serves two principal purposes. First, it may
be part of the selective process in mating, as when male birds
strut their plumage in the nuptial ceremony, and as when stags
lock their horns in mortal combat in competition for a doe. Sec-
ond, among some animals that inhabit distinct territories, as for
example lions, or baboons living in a forest, the exaggerated size
and fighting equipment of the males permit them to serve the
function of a border patrol in human communities. The male
keeps rivals off his feeding ground and away from his wife or
wives. Neither the male lion nor the male baboon is any better at
obtaining food than his womenfolk; in fact, among lions the fe-
male excels at hunting. These animals expend their biological
capital for territorial defense, just as we spend the bulk of our tax
money for atomic submarines and missiles.
In fossil man there is evidence of sexual dimorphism, but it is
clouded by the paucity of material available for study. In living
races a great variability can be seen. Australian aborigines and
western Europeans are highly variable; Mongoloids little. As Ti-
betans dress and wear their hair alike, it is sometimes difficult to
27
How Species Have Evolved
tell whether any one person is a man or a woman. This does not
mean that sexual dimorphism is the same as pedomorphy, for
some populations with little sexual dimorphism are in certain
ways gerontomorphic. No one could call a Plains Indian infan-
tile, and his women can be huge and craggy. It is difficult, then, to
decide whether certain racial traits, like the absence of a beard in
many Mongoloid males, are the result of pedomorphy, of a lack of
sexual dimorphism, or of some other aspect of the endocrine story
yet to be discovered.
In any case, the presence or absence of marked sexual dimor-
phism is an inherited racial trait that distinguishes some living
populations from others. This trait may date back to remote an-
tiquity since it was not involved in the complex of evolutionary
changes that led from Homo erectus to Homo sapiens. Of this we
may be fairly confident because the two races that have achieved
the greatest cultural advancement, the Caucasoid and the Mongo-
loid, stand at opposite poles in this respect. At the other end of
the cultural scale, so do the Australian aborigines, who show
marked sexual dimorphism, and the African Bushmen, who show
little of it.
How Species Have Evolved
Like all men, all species must eventually die. Just as
some men perish with neither issue nor close kin and others
achieve partial immortality through the transmission of some of
their genes to their offspring, or more remotely, by the survival
and reproduction of their brothers and sisters — so some species
become utterly extinct whereas others live on, in a shadowy way,
through one or both of two evolutionary mechanisms, succession
and branching. Succession is also called phijletic evolution or
anagenesis; the technical word for branching is kladogenesis.
Evolution through succession occurs when a genetically iso-
lated population acquires a new and favorable hereditary trait
that is controlled by a single gene or by a complex of genes op-
erating in concert. Then the new trait gradually replaces the old
one through natural selection.
2 ^ The Problem of Racial Origins
Evolution through branching occurs when two or more geo-
graphically separate populations of a single, polytypic species be-
come genetically isolated from one another and then evolve into
species of their own.
Succession tends to favor a process known as general adapta-
tion whereas branching works rather through special adaptation,
but the two are not mutually exclusive.
General adaptation involves the acquisition of a new trait or
trait complex that is useful in more than one environment and un-
der various different circumstances. Warm-bloodedness in birds
and mammals is one example. Another is an increasing intelli-
gence, which many forms of animal life have developed through-
out geological history. A more limited example is the power of
speech, which is useful to all men.
Special adaptation involves the acquisition of a new trait or
trait complex that is useful in a single environment under special
circumstances. It is the process which enables an animal to resist
heat, cold, or bright light, to see well in dim light, to run faster or
to swim better than its fellows, or to live without water in deserts,
and which gives it many other such specializations. Special adap-
tation led the ancestors of the whales from the land back into the
sea, and general adaptation gave them the intelligence needed to
communicate with one another, by a system similar to sonar, and
to survive, as mammalian populations, in their aqueous medium.
General adaptation tends to lead a species into evolution by
succession because most species are polytypic, and a polytypic
species includes several populations living in different environ-
ments. Each of these populations becomes adapted to its special
environment to a certain degree, but it cannot speciate by branch-
ing as long as it remains in genetic contact with its sister popula-
tions, since new traits involved in local specialization cannot com-
pletely replace old ones while genes continue to flow back and
forth. If, however, in one or more populations a new trait appears
which is equally favorable to all the populations and in all the en-
vironments occupied by the species, then the existing gene flow
will help the new trait replace its predecessor in all the compo-
nent populations, including that or those in which it started. By
this process the old species evolves as a unit into a new species. At
How Species Have Evolved 29
the same time speciation need not prevent the component popu-
lations from carrying their old, partial specializations, such as to
heat and cold, from one species into another.
If, however, a single population of a polytypic species becomes
physically isolated from its fellows, so that gene flow is completely
interrupted, then that population can evolve by branching. Now
special traits that have no general value can completely replace
the old ones that used to flow in over the border. If such a popula-
Fig. 1 How One Polytypic Species
Can Evolve Into Another. Above:
Five subspecies, in peripheral contact
with each other, are illustrated by five
circles, numbered 1 through 5. A
mutation favorable to all five arises in
No. 3. It spreads to Nos. 2 and 4, and
is carried by further peripheral gene
flow to Nos. 1 and 5. When all five
subspecies have it, the species has
begun to evolve into a new one by
anagenesis — evolution through succes-
sion. Below: In this example the fa-
vorable mutation arises independently
in Nos. 3 and 5, and, except for the
direction of gene flow between Nos. 4
and 5, speciation takes place as in the
first example.
tion happens to be confined to a small space, such as an island,
and has no natural enemies, it can become a monotypic species as
specialized as the dodo, the classic example of this process.
Although the component populations of a polytypic species
evolve as a unit, they cannot do so simultaneously since it takes
time for a mutation to spread from one population to another. If
we measure time on the broad scale of tens of millions of years
used by paleontologists, these changes may appear simultaneous.
3° The Problem of Racial Origins
but if we measure it on the geologically microscopic scale of the
last 700,000 years, which is the age of man, we will see that
related populations, which in our case are subspecies, passed from
species A, which is Homo erectus, to species B, Homo sapiens, at
different times, and the time at which each one crossed the line
depended on who got the new trait first, who lived next to whom,
and the rates of gene flow between neighboring populations.
Whether a new species is polytypic or monotypic, whenever it
arises the evolutionary process is essentially the same. The new,
critical trait responsible for speciation first appears in a few indi-
viduals, and its presence makes little difference to the population
in which it arises. It may even appear and disappear several times
before it takes hold. But after it has begun to spread, a point is
reached when those who have it begin to outnumber those who
don t. This point is marked by a rapid growth in population. The
particular population has gained an advantage over competing
species in its own lebensraum, and in the process it has become a
new species of its own.
It need not, however, have completely lost the gene or genes for
the old trait that is on the way out. After the new species has es-
tablished itself, become stable in numbers, and reached a new
equilibrium with the other species of plants and animals in its en-
vironment, the old trait may completely disappear. At that point a
second and final threshold of speciation has been crossed. One
may say that a new species has come into existence when it has
acquired a new and more favorable ecological position, and that
it has reached maturity when the traits responsible for these
changes have completely replaced their predecessors. By the time
the second threshold has been crossed, as likely as not a new spe-
cies-forming mutation shall have begun to appear, and the cycle
has started over again.
It is easy to understand, then, why some populations within any
polytypic species have come closer, at any given time, to the sec-
ond threshold of speciation than other populations. In man some
groups of people alive today have preserved archaic traits, diag-
nostic of Homo erectus, in a higher percentage of individuals than
other populations. For example, more natives of New Caledonia
3i
How Species Have Evolved
have big teeth and heavy brow ridges than a corresponding per-
centage of Japanese.
This and similar disparities can be explained in two ways.
( 1 ) The more archaic population acquired the new trait complex
that led to speciation later than the more modern population did.
(2) After crossing the first threshold of speciation, the more ar-
chaic population has been discarding its old traits at a slower rate
than the more modern population.
Both explanations can be true at the same time. There is no
necessary correlation between the time at which a threshold was
crossed and the rate of change that follows the crossing. In either
case, the critical mutation may have been original to the popula-
tion concerned, or it may have been acquired by gene flow from a
neighboring population. The older the trait the more likely that it
was original; the younger the trait the more likely that it was de-
rived from outside.
In any event, once a species has come into being, the old spe-
cies from which it evolved is extinct. There are several kinds of ex-
tinction: utter extinction without issue, which is commonest
among monotypic species; extinction through absorption, by
which a subspecies ceases to exist as a separate entity when its re-
maining members are taken into the body of another; and extinc-
tion through successive evolution, which is the process we have
just described.7
In the case of man, only the second and the third kinds of ex-
tinction can be traced. The Tasmanian aborigines who died out
in the nineteenth century have living survivors among the racially
mixed inhabitants of the islands between Tasmania and Austra-
lia, and the Fuegian Indians of South America are disappearing
into the mixed population of that continent. But neither Tasma-
nians nor Fuegians were whole subspecies. The Australoid and
Mongoloid divisions of man to which they belong survive in large
numbers elsewhere. There are also, in a sense, degrees of extinc-
tion, for it takes a long time, on our human time scale, for one
7 E. H. Colbert: “Some Paleontological Principles Significant in Human Evo-
lution,” in W. W. Howells, ed.: Early Man in the Far East (Philadelphia: Am.
Assn. Phys. Anth.; 1949), pp. 103-47.
32 The Problem of Racial Origins
species to replace another completely, and in that sense some hu-
man races are more nearly extinct than others.
On the Life Spans of Mammalian Species 8
Although the antiquity of Homo sapiens will be the subject
of detailed study in later chapters, we may here profit from a con-
sideration of the life spans of our fellow mammals during the
Pleistocene and Recent (or post-Pleistocene) periods, the only
periods in which man or any of his close kin are known to have
lived. By international agreement the beginning of the Pleisto-
cene has been established at the point when modern genera of
elephants, horses, oxen, deer, and some other large mammals were
first seen on the continents of the Old World, excluding Australia.
The movement that brought them in, mostly from the New World,
took place about one million years ago.
Before that stretched the vast temporal expanse of the Tertiary
— Paleocene, Eocene, Oligocene, Miocene, and Pliocene— com-
prising some 77 million years, the Pliocene alone taking up some
12 million. During this long span individual species were born,
flowered, and died at what seems to us a leisurely pace. The life
expectancy of a mammalian species was then anywhere from one
to eight million years.
During the first 300,000 or 400,000 years of the Pleistocene this
pace continued, but it was suddenly quickened in various parts
of the Old World, particularly its northern portions, by geological
events. The planet’s crust wrinkled more rapidly than before, rais-
ing the toothed edges of mountain ranges and creating great con-
trasts of climate, both regional and seasonal. First mountain gla-
ciers, then continental icecaps crawled forth and melted away,
blowing, like pairs of bellows, alternately cold and warm. In large
tropical land masses, as in much of Africa, the bellows blew wet
and dry.
8 This section is based on many sources, including books by G. G. Simpson.
However, specific facts and figures come principally from two works of Bjorn
Kurten: “Rates of Evolution in Fossil Mammals,” CSHS, Vol. 24 (1959), pp.
205-i5, and “Chronology and Faunal Evolution of the Earlier European Glacia-
tions, SSF-CB, Vol. 31, No. 5 ( i960) pp. 1—62.
33
On the Life Spans of Mammalian Species
In response to these changes, new species evolved rapidly.
Many became extinct, but others survived. The life expectancy
of a species now dropped to a mere 360,000 years. At a point in
time pegged at 300,000 years ago, all or nearly all the living mam-
mals of the European and neighboring fauna, which were fox-
sized or larger, had come into existence. The species which have
since appeared are bats, insectivores, and rodents, all small ani-
mals. During the last 75,000 years, no new mammalian species
seem to have evolved at all. Three hundred thousand years ago
the evolution of new species of medium-sized and large mammals
came to a halt. The heyday of speciation was over.
The oldest known Homo erect us is believed to be 700,000 years
old. He appeared during the period of frenzied mammalian spe-
ciation mentioned above, and seems to have lasted until less than
100,000 years ago in remote parts of the Old World. His known
life span as a species, about 600,000 years, was within the normal
range for a mammal of his size and vintage. As I shall show in
Chapter 11, Homo sapiens appeared about 250,000 years ago in
an archaic form. Completely modern forms of our species ap-
peared at least 35,000 years ago. Unless our species is a curious
exception to the rules by which the game of speciation is played,
Homo sapiens should go back to 360,000 or 300,000 years ago.
This figure would place Homo sapiens in the fauna to which he
belonged, and would give Homo erectus, who appeared exactly
when he should have, ample time for speciation by succession.
So much for the actuarial statistics of Pleistocene and Recent
species. With subspecies the reckoning is more difficult because
subspecies are not easy to sort out when found among fossils. We
have no satisfactory information except that subspecies of the ibex
have been traced back at least 230,000 years.9 In the case of man,
the subspecies of Homo sapiens are probably of different ages, de-
pending on the times at which regional populations of Homo
erectus, in one way or another, crossed the sapiens threshold. But
all of them did this before the end of the Pleistocene.
In modern times we have seen whole tribes and peoples disap-
pear after their lands had been invaded by Europeans and other
9F. Zeuner: Dating the Past (London: Methuen & Co.; 1952), pp. 383-4.
34
The Problem of Racial Origins
culturally dominant strangers. The native Tasmanians are gone,
and so aie the Indians of Lower California. The Andamanese of
the main islands, the Fuegians, and many others are on their way
out. These sad cases of ethnic oblivion give us a feeling that hu-
man history is a long record of utter extinctions, but this is not
true.
All species are destined to become extinct, but, except as they
are parts of species, subspecies need not follow this rule. By defi-
nition, species do not ordinarily interbreed, but subspecies do.
The Tasmanians were absorbed by the Caucasoids who replaced
them on their island. A mixed Tasmanian-European population
survives today. If the Indians of Lower California left no mixed
descendants— which is unlikely— other Indians very much like
them are still alive. When subspecies disappear, they usually, if
not always, do so by absorption. Their genes linger on polymor-
phously with those of their conquerors, to re-emerge, now and
then, when needed. The principle is that when a population has
been invaded by members of another race the genes that give it
its special adaptation to its local environment retain their selective
advantage and eventually come to characterize the mixed popu-
lation through the process of natural selection. For example, cen-
tral Europe was invaded from the East many times from the Neo-
lithic through the Iron Age, but central Europeans still look more
like the hunters of the Mesolithic than like the invaders.1 Without
the concepts of absorption and re-emergence it would be difficult
for us to explain the physical diversity and geographical distribu-
tion of the living human races.
Part of this diversity may be relatively new. I refer here espe-
cially to the reduction in body size that has affected many species
of mammals since the end of the Pleistocene, some 10,000 years
ago. As will be explained in Chapter 3, extreme cases of size re-
duction in plants and animals take the form of dwarfing, which
means that an irreversible genetic change has taken place. Our
species includes a dozen or more populations of dwarfs, living in
Africa, southern Asia, and Indonesia. As far as we know, all hu-
man dwarf populations are geologically recent.
1C. S. Coon: The Races of Europe (New York: The Macmillan Co.; 1939).
Genetic Principles and the Origins of Races
35
Genetic Principles and the Origins of Races
In recent decades the pursuit of anthropometry has de-
clined, except for applied anthropology. Instead of measuring the
bodies of the last remnants of aboriginal populations, anthro-
pometrists measure military personnel and civilians in order to de-
sign railroad and airplane seats and space suits. Doctors of Phi-
losophy have become tailors to the new age of science. On the
other hand, the pursuit of human genetics has become popular,
paiticularly the study of the frequency in populations of blood-
gioup genes, taste thresholds, mid-digital hair, and hairy ears.
In tracking down the lines of descent of fossil men, none of
these characteristics is useful. Thieme and others have shown that
it is impossible, using present techniques, to determine the blood
type of samples of bone, for they all tend to absorb a group A sub-
stance fiom the ground." Dead men cannot taste noxious chemi-
cals, and the hair on their fingers and ears has long since de-
cayed. What could be done, however, is to work out the relation-
ships between fossil specimens and populations in terms of details
of tooth structure, for teeth do not change with age except to be
worn down. Molar cusp-numbers, the presence or absence of a
kind of curvature of the incisors known as shoveling, and many
other features that are preserved in the fossil record are just as
useful for genetic studies as blood groups are among the living,
and paleontologists have long relied on teeth. Although much
work has been done on human teeth no one has yet produced a
work of synthesis covering all fossil specimens by means of which
they could be compared with living populations.
Limited as the direct application of genetics is in the study of
fossil man, the theoretical aspects of that science have helped us
greatly. They have taught us that the unit of inheritance is neither
the individual nor the arbitrarily chosen type, often identified
with an individual, like Nordic, Dinaric, Neanderthal, and Cro-
Magnon, but the population, and that each population has its pool
2 F- P- Thieme and C. M. Otten: “The Unreliability of Blood Typing Ancient
Bone,” AJPA, Vol. 15, No. 3 (1957), pp. 387-97.
36 The Problem of Racial Origins
of genes with several possible alternates, known as alleles, from
many if not all loci. We also know that because individual muta-
tions recur at characteristic rates, resemblances between popula-
tions of the same species do not necessarily imply recent common
descent. All curly-haired populations do not have to be descended
from a common curly-haired ancestor. Pockets of blondism found
among nonwhites need not be explained by Viking invasions, nor
all Pygmies be considered as having derived from a single tribe.
An acquaintance with the principles of genetics may also help
us solve the central problem of this book— that is, to discover
how long ago the ancestors of the human subspecies parted com-
pany. We have learned, for example, that evolution proceeds trait
by trait, one mutation, recombination, or whatever, at a time. If
parallel mutations have been occuring in two populations, we
cannot expect a large number of identical changes to have taken
place at once in each group. Changes in the skeletons of fossil
men from period to period in each major area seem to have in-
volved very few factors, not many of them visible below the neck.
Brains have grown larger and brow ridges smaller. Jaws have
sprouted chins and teeth have grown smaller in various degrees.
Whole sets of these changes can be linked together as common
products of one or more shifts in endocrine balance, shifts advan-
tageous in an increasingly group-oriented society in which self-
control comes to be more conducive to survival than a hot temper.
Other changes may simply reflect a reduction in chewing, espe-
cially after the invention of cooking. If in each of several related
populations, living in its own territory, changes like these took
place not all at once but in sequence, it is possible that each sin-
gle, parallel mutation prepared the ground for the selective ad-
vantage of the one that followed it.
On the other hand, if these sequences of genetic change were
initiated in some of the populations by sexual contacts with peo-
ple from other regions (peripheral gene flow), it would be diffi-
cult for us to detect this outside influence from an examination of
the skeletons of the resulting mixed population because other
genes transferred by the same contact might be disadvantageous
in that particular area and would have been eliminated by natural
selection.
Genetic Principles and the Origins of Races 37
Although we cannot hope to settle the question of parallel
evolution versus peripheral gene flow in the evolution of each race
by examining fossil bones and nothing else, such a study may
show us how far back in time the various geographical races go.
Some of our subspecies are characterized by traits that seem to
have had little relation to either climate or culture during the
known history of man, and whatever selective advantages or dis-
advantages they may have must have been acquired long ago.
Among these traits are the architecture of the teeth, the shape of
the nasal bones, and the degree of flatness of the face. If various
combinations of these traits can be seen to have persisted in their
special geographical regions despite other changes of a more
clearly phyletic evolutionary nature, then the antiquity of indi-
vidual races may be established. In any case, no form of evidence
is unwelcome and only by a close study of detail can we hope to
solve this and related problems.
8
2
K
K
K
EVOLUTION THROUGH
ENVIRONMENTAL ADAPTATION
In this chapter we shall discuss the effects of size, space,
numbers, and climate on the direction and rate of evolution.
Other principles of change will be examined, in addition to those
mentioned in the first chapter. By comparing man with other ani-
mals we shall see, in particular, how adaptation to the external,
nonhuman environment helped shape the living races of man into
their present forms.
Body Size, Food, Space, and Climate
At some turning point in the evolution of our ancestors men
became hunters. Instead of relying on roots, fruits, and small,
slow-moving animals for their food, they began to compete suc-
cessfully with the great carnivores and could feed themselves and
their families wherever meat was to be found.
Among other carnivores a natural relationship exists between
the size of the predator and that of his prey. A fox cannot kill a
zebra, but a zebra is the favorite food of lions. Properly armed,
however, a man can kill an animal of any size. As he began to do
so, it was probably advantageous, all else being equal, for him to
be large as well as muscular. At any rate, we have indirect evi-
dence that our ancestors grew larger at about that time. This
placed them in the elite company of large land mammals and sub-
Body Size, Food, Space, and Climate 39
jected them to some of the special evolutionary rules that govern
such species.
One is that they are few. At present there may not be more than
sixty other species of our size or larger,1 out of 3,500 species in the
class of mammals. A second is that most of these have only one
species to a genus. This ratio is characteristic of the other large
primates — the orang, the chimpanzee, and the gorilla — as well
as of the elephant, the hippopotamus, the rhinoceros, and the
bear. The larger these animals the more likely that they will be
the only species in their genus, at least at any one time. In his
utilization of terrain man is in a class with the largest mammals of
all; primitive hunters destroy more forest by burning than ele-
phants do by uprooting trees. If, after the beginning of hunting,
the genus Homo ever consisted of more than one species at a time,
not counting species in the act of transition, as from Homo erec-
tus to Homo sapiens, he would have constituted a curious excep-
tion to a well-established rule.2
A third rule is that individual animals of large size take up dis-
proportionately more room than small animals, and that the rela-
tionship between the sizes of their ranges is believed to be loga-
rithmic.3 A large animal needs a great deal of space not only for
feeding and drinking but also for concealment in the heat of the
day and for sleeping and reproduction.
From the time that man became a hunter, if not before, he was
a social animal living in groups of families with an optimum popu-
lation somewhere between twenty and forty persons.4 As the area
required per person must be multiplied by the total in the group,
the territory of a feeding unit of this size had to be considerable.
Once the ability to eat meat had extended the potential range of
man as a species to the limits of the continental land masses of the
Old and New Worlds, nothing could stop him from filling these
1 The uncertainty is due mainly to a lack of agreement on the classification of
the Bovidae — cattle, sheep, goats, antelope, etc.
2E. Mayr: “Taxonomic Categories in Fossil Hominids,” CSHS, Vol. 15 (1951),
pp. 109-18.
3 G. E. Hutchinson and R. H. MacArthur: “A Theoretical Ecological Model of
Size Distributions among Species of Mammals,” AN, Vol. 93, No. 869 (1959), pp.
117-25.
4 The basis of this calculation will be stated later.
40 Evolution through Environmental Adaptation
spaces except natural barriers, such as glaciated mountains,
bodies of water, empty deserts, and extremes of environment to
which he was not accustomed, such as great cold, heat, drought,
and high altitude.
Some of these extremes he overcame by cultural means, par-
ticularly after he had acquired fire. His ability to invent and make
adequate housing, clothing, and containers as well as effective
weapons must have placed a premium on the kind of intelligence
that governs these capacities. In every region where, in addition
to hunting, environmental problems had to be mastered by tech-
nology, parallel evolution in the direction of higher intelligence
must have been in operation.
Once man s inventive genius had made it possible for him to
live in extreme environments previously barred to him, a new
burden was placed on his physiology because he could not, with
his incipient skills, overcome all climatic obstacles. We must ex-
pect to see the results of genetic responses, through natural selec-
tion, to differences in environment, and we must know how to in-
terpret them, for the patterns they take will tell us much about the
early history of our genus and species.
If such differences occur and the subspecies of man turn out to
be clearly divided on this basis, so that one is adapted for wet
heat, another for dry heat, a third for cold, a fourth for high alti-
tude, and so on, then it will be likely that at the time of the dis-
persal of the subspecies the ancestors of all of them belonged to a
single, monotypic population. If, on the other hand, the existing
subspecies do not entirely fit this scheme, and if in addition cer-
tain subspecies include regional populations adapted for different
climates, then it will appear that Homo has been polytypic for a
very long time and that the dispersal of our ancestors into differ-
ent regions took place very early, before the beginning of man’s
career as a hunter and before he possessed the cultural means to
invade and inhabit the regions of the earth s surface previously
unavailable to him.
In pursuing this inquiry we shall not be exploring virgin terri-
tory. Zoologists have been faced with similar problems for over a
hundred years. What happened to human beings once they had
come to live in diverse environments had happened to other ani-
The Face of the Earth 41
mal species many times before. The study of animal evolution
through environmental adaptation is a part of zoogeography,5 a
well-documented scientific discipline more than a hundred years
old. By reviewing its major principles and studying some of the
data it has uncovered we may determine what to look for in man.
The Face of the Earth
As our telescopes improve and we learn more and more
about the surfaces of other planets — most of which are excessively
hot or cold, or vary from one extreme to the other between night
and day, and lack the friendly mists and rain that water our
woods and fields — the more we appreciate the infinite variety and
manifold advantages, to creatures like us, of the face of the earth,
our home.
Far from being simply a playground to run about, sleep, feed,
and breed in, the skin of our planet, with its myriad variations,
has been a major determinant in the evolutionary process. Evolu-
tion, itself a product of variability and change, has been cumula-
tive, keeping pace through geological time with the ever increas-
ing rate of differentiation of the surface features of the earth. The
planet s crust has wrinkled faster, in cooling, than wind, rain,
snow, ice, and all the other forces of erosion have been able to
wash away, grind down, flatten, and otherwise homogenize it. The
same forces have made the products of evolution increasingly
complex and heterogeneous, increasingly sensitive, and increas-
ingly aware of themselves and of their surroundings.
Eight factors have affected the face of the earth from the zoo-
geographic point of view: the clockwise rotation of our planet,
which creates westerly winds; the zonal differentation of the
earth, which makes some latitudes cold and others warm; the tilt
of the earth’s axis, which creates seasons; the relative sizes of land
masses, which emphasize or diffuse seasonal change and give
populations breeding grounds of different magnitudes; the rise of
5 The study of plant distribution is called phytogeography. Phytogeography +
zoogeography = biogeography, the study of the distribution of living things. Bio-
geography is a unit; its two components are interdependent.
42 Evolution through Environmental Adaptation
mountain ranges, which permit altitude to substitute for latitude
as a climate maker; the bodies of salt water, which prevent ter-
restrial animals from crossing from one land mass to another; and
the land bridges and strings of islands, which allow certain quali-
fied animals to filter through. The eighth factor is time, particu-
larly the last million years, which have seen the icecaps of the
world alternately crawl forth and shrink back three times, with
consequent stress and displacement of many forms of animal life,
including that far-ranging genus, Homo.
Land Masses 6
According to standard school geographies, the world con-
tains seven continents, but for present purposes these will be con-
sidered as five. Antarctica can be disregarded because it is un-
inhabited. Europe is not a real continent; the Greeks distin-
guished their own peninsula from Asia, which lay on the other
side of the Aegean, and this split has since been carried north past
the Caspian Sea barrier onto the steppes of Russia. Europe is a
highly favored peninsula of Asia. We shall call the combination
Eurasia, as do most geographers.
Of the five continents remaining — Eurasia, Africa, North
America, South America, and Australia — the first four are strung
together in one fashion or another. The Isthmus of Suez ties Af-
rica to Eurasia as it has done for a long time. Both the Bab el
Mandeb and the Straits of Gibraltar are deep but narrow salt-
water channels, and botli were open and ice-free throughout the
Pleistocene. Eurasia, the greater and more varied of the two seg-
ments, is nearly twice as large as Africa, with 21 to the latter’s 12
million square miles, and it contains bits and pieces of all the cli-
mates of the world, whereas Africa lacks mid-latitude forests, bo-
real forests, and taiga.7
G In this and the following two sections of Chapter 2, I am drawing heavily
on P. J. Darlington, Jr.: Zoogeography (New York: John Wiley & Sons; 1957).
For the Pleistocene, the best source is J. K. Charlesworth : The Quaternary Era,
2 vols. (London: Edward Arnold; 1957).
7 These terms are taken from Preston James’s Outline of Geography (Boston:
Ginn and Co.; 1935).
Land Masses
43
Eurasia and Africa are tied together at a latitude of 30° N., and
as Suez is normally frost-free, it is not too cold at sea level for
most forms of terrestrial life to traverse. Africa is mostly a plateau
bent around a cup of low-lying equatorial rain forest; and from
the edge of the Sahara to the Cape of Good Hope there stretches
an essentially homogeneous environment of grasslands, savannas,
and seasonal forests high enough so that temperatures vary little
from one latitude to another and grazed throughout by more or
less the same kinds of animal herds. By contrast, Eurasia is built
like a tent with a pole in the middle and drooping sides. The lofty
land mass of Tibet brings an approximation of arctic conditions
to a large area partly located in the same latitude zone as Suez. It
also partitions off much of southern Asia. The line of mountains
reaching diagonally across the map from the Tian Shan to the Ber-
ing Strait cuts the northern half of the continent into a north-
western and a far-eastern segment.
The erstwhile land bridge across the Bering Strait, which con-
nected Eurasia with North America, was a broad, flat, ice-free
highway that appeared during periods of glaciation whenever the
ocean level was lowered by the immobilization of water in the
form of ice at the poles. The land bridge last appeared probably
between 70,000 and 8,000 b.c., either during this entire period or
in parts of it. Although it lay at an altitude of 66° N., the south-
ern shore of the bridge may have had mild winters at this time,
being protected from the arctic waters and tempered by the west-
ward flow of the Japanese current. Animals able to live through a
moderately cold winter could have crossed the bridge in either
direction, and many of them did.
North America, with 8.3 million square miles, is smaller than
Eurasia or Africa and differs from both in land formation.
Whereas Africa is predominantly a plateau and Eurasia a ring of
subcontinents with most of its mountains running east and west,
in North America the western and eastern ranges run north and
south, leaving a wide trough in the middle which creates extremes
of climate at many widely separated points, so that one can shiver
in Houston in winter and swelter in summer in Saskatoon.
South America, with 6.8 million square miles, has been con-
nected to North America by the Isthmus of Panama since the be-
44 Evolution through Environmental Adaptation
ginning of the Pleistocene a million years ago, but during the en-
tire 60 million years of the preceding Tertiary it was isolated by
salt water. Like Africa, it has a plateau running across the equa-
tor, and this plateau is as high as the Tibetan one although its sur-
rounding peaks are a little lower. But it is narrower, and the
equatorial rain forest it shelters is, at the present geological mo-
ment, the world’s largest.
However, if we return in time to the last glacial advance, and in
space to southeast Asia (see Map 2), we see that a vast area of
some 800,000 to a million square miles, known as the Sunda Shelf,
was then incorporated, as geologists believe, onto Indochina, Ma-
laya, and the islands of Indonesia east of the Bali and Macassar
Straits. If, as may be presumed, this lowland was largely covered
with rain forest — for it was a wet period — it may well have been
as large as the South American rain forest, or even larger. A rain
forest of this size is a fertile breeding ground.
Five hundred miles to the south and east lay a continental area
of nearly four million square miles, including the present Austra-
lia (3 million square miles), Tasmania, New Guinea (300,000
square miles), and some of the Melanesian Islands, joined by an-
other now-submerged stretch of lowland, the so-called Sahul
Shelf (over 580,000 square miles). When the sea rose at the end
of the Pleistocene, this land mass was split into its present com-
ponents.
Zoologically these now separated regions are still a unit. An-
thropologically we can likewise consider New Guinea, Tasmania,
and some of the Melanesian Islands as recently separated periph-
eries of a fair-sized continent the center of which is the Australian
desert. More specifically, woolly hair is characteristic of the
Papuans, Tasmanians, and a few of the coastal Australian aborigi-
nes. Most of the Australians have straight or wavy hair. Woolly
hair, therefore, is geographically peripheral to straight hair in what
is left of the former, larger continent.
The significance of the Sunda and Sahul shelves is clear. In no
part of the world other than southeast Asia, Indonesia, and Aus-
tralia are the seas so shallow that vast interconnecting land masses
could have been created when the icecaps of the polar regions
trapped enough water to lower the sea levels in many parts of
THALIA
46 Evolution through Environmental Adaptation
the world by forty fathoms below their present shorelines. The
Snnda and Sahul shelves are the only real “lost continents.” Not
only did they join lands now separate, but they may also have
served as bellows to suck in and blow out early human popula-
tions.
Two important facts emerge from this survey of global land
masses. The Northern Hemisphere is the land hemisphere, and
the Southern the realm of ocean. Therefore the land masses situ-
ated in the north are more continental in climate, that is, more
extreme in seasonal change, and stormier than the southern lands,
where less meteorological change is taking place. The Old World
with its combined mass of Eurasia and Africa, which are divided
only by narrow seas, is a huge and varied breeding ground com-
pared to the New with its smaller masses of North and South
America, which meet effectively at a single point only. One would
expect more to have happened biologically to land animals in the
Northern than in the Southern Hemisphere, and also in the Old
World than in the New. These expectations have been fulfilled,
particularly in the case of man.
Barriers and Breeding Areas
The most conspicuous barriers in the world are probably
mountains, especially such lofty breath-takers as the Himalayas,
but even their rims can be crossed, by animals as well as people’
and the principal hindrance they offer is the rapid temperature
giadient rather than the steepness of terrain. Deserts, too, are
baniers; there lack of moisture, more than temperature, does the
screening, and except in sandy stretches the terrain itself offers
little impediment to travel.
The greatest barriers of all, however, are stretches of salt water.
That is why the Azores, when first occupied in the fifteenth cen-
tury, had no land animals except birds and a local lizard, and why
the Australian continent contained aboriginally no placental
mammals except man, the rat and dog, which went with him,
and the air-borne bat. That is why South America contained an
almost unique vertebrate fauna when the North American animals
Genetic Drift 47
began infiltrating over the newly formed Panama bridge at the
end of the Pliocene. Lesser barriers such as mountains and des-
erts serve as screens rather than as roadblocks. While holding
back most species, they let dominant ones through to take over
new territories. In the case of subspecies, especially qualified in-
dividuals can get from one breeding ground to another and
spread their genes in the new population.
It is a general rule that relatively numerous populations living
in large breeding grounds tend to be dominant over others that
have lived in smaller areas. The larger the number of animals in a
population, the greater the mathematical chance they have of un-
dergoing a rare, favorable mutation that can spread through-
out the group by means of natural selection. Since in small, iso-
lated populations there are fewer individuals there are also fewer
mutations, too few in some cases to include any of the uncommon,
favorable ones. At the same time, owing to lack of competition in
a sheltered environment, some of the commoner, unfavorable mu-
tations can spread unhindered through such a small, sheltered
population and eventually bring about deterioration or even ex-
tinction. That is why islands are being constantly repopulated by
stray sets of dominant species that happen to drift or be blown in
from continental land masses.
Genetic Drift
Nevertheless, all mutations need not be perceptibly or
measurably favorable or unfavorable in any given situation. In
Europe, for example, it can make no conceivable difference to a
man’s chances of survival and reproduction whether his hair is
straight or slightly wavy. In a large population, a neutral or in-
different mutation will not ordinarily spread rapidly, nor will it
necessarily be lost. It can be expected, all else being equal, to
maintain a low frequency in a large gene pool. In a small popula-
tion, on the other hand, it can easily be lost through sheer chance
— if, for example, the three persons out of ten who have it are
eaten by a tiger. The mutation could also spread through the
same small breeding unit if the tiger ate the people who did not
48 Evolution through Environmental Adaptation
have it instead of the others. Gene frequencies, then, change more
rapidly in small than in large populations. The process by which
such fortuitous changes become major characteristics of popula-
tions is called genetic drift, or the Sewall Wright effect, after its
discoverer.8
Once genetic drift has taken place, the chances are that the
population in which it has occured will become extinct, because:
( 1 ) the reduction in population which permitted the drift may
also have reduced the total number of breeding individuals below
the safety level needed for survival; and (2) few genes chosen by
chance are likely to be superior to their alternate alleles from the
standpoint of survival.
If the population survives and multiplies, this may be because
the genetic characteristic or characteristics chosen by drift were
favorable for survival in the first place, and the drift merely sped
up the process of selection. In the long run, the frequency of this
gene or of these genes in the pool would have risen to an optimum
level in any case, without danger of extinction.
Genetic drift is often invoked to explain differences between
species and subspecies in characteristics that are of no detectable
value in natural selection. As our knowledge of genetic processes
grows and as our ability to detect selective values increases, we
need this theory less and less.
The Dominance of Groups
Dominance has two meanings in zoology: the dominance of in-
dividuals in social groups, as shown by the peck order and the
like, and the dominance of one kind of animal over another. We
are concerned here with the second meaning only.
Groups of dominant animals may range in diversity from whole
orders to families to genera and even to species. Examples are
the carps (family Cyprinidae ); the common frogs (genus Rana );
the common snakes (family Colubridae) ; the perching birds
8 S. Wright: ‘On the Role of Directed and Random Changes in Gene Fre-
quency in the Genetics of Populations,” Evolution, Vol. 2, No. 4, (1948), pp.
279-94-
49
The Dominance of Groups
(order Passer es ); the rats and mice (family Muridae ); and the
human species ( Homo sapiens). Even within a species such as
ours, certain subspecies and races may show dominance over
others. This is part of the evolutionary process.
Dominant groups result from a combination of factors that
render them particularly successful in withstanding climatic
stress, especially cold; in finding and utilizing food; and in re-
producing efficiently under varying circumstances. The perching
birds, for example, achieve these ends partly by migration; the
rats and mice by storing food and by burrowing underground to
escape predators and the rigors of the weather. In the case of
man, he is capable not only of using fire and tools intelligently in
organized social units but also of undergoing a certain amount of
physical adaptation to certain environments.
As a rule, the breeding grounds of dominant animal groups are
situated in the centers of the land masses they occupy, with the
result that the animals are forced into competition for their eco-
logical niches by rivals from all sides. If in addition to being cen-
trally located, the breeding grounds are in cool regions, then the
species living there will produce more offspring than the same or
corresponding species in the tropics,9 not because of greater fer-
tility, but because in warm regions many fetuses are lost through
the failure of sufficient pituitary hormone ACTH to reach the
embryo from the mother. As this hormone normally balances the
adrenal cortisone, which has no difficulty getting through, an ex-
cess of cortisone causes the resorptions.1
This and other observations partially explain why the cooler
portions of the Old World had fewer species, but larger popula-
tions, than its tropical regions, and why some of these populations
reinvaded the tropics, with varied success.
Zoogeography can also explain many instances in which groups
9B. Rensch: “Some Problems of Geographical Variation and Species Forma-
tion,” PLSL, 149th session (1936-7), pp. 275-85. Also Homo Sapiens, vom Tier
zum Halbgott (Gottingen: Vandenhoeck and Ruprecht; 1959).
1 W. V. Macfarlane, P. R. Pennycuik, and E. Thrift: “Resorption and Loss of
Fetuses in Rats Living at 350 C.” J. Physiol., Vol. 135, No. 3 (1957), pp. 451-9.
Also S. Brody, A. C. Ragsdale, R. G. Yeck, and D. Worstell: “Milk Production,
Feed and Water Consumption, and Body Weight of Jersey and Holstein Cows
in Relation to Several Diurnal Temperature Rhythms,” RBMO, Vol. 578 (1955),
pp. 1-26.
50 Evolution through Environmental Adaptation
failed to acquire dominance. Animals that inhabit peripheral
shores or small islands lead sheltered lives and may develop local
peculiarities without facing the pruning effect of rivalry. That is
why early mariners found dodo birds strutting around Mauritius
and giant tortoises ambling over the glades of the Galapagos.
These facts, incidentally, were not lost on the youthful Darwin
who voyaged on the Beagle. That is also why rabbits and foxes,
when let loose in Australia, raised such havoc with the local fauna,
and why, in another sense but following the same principle, the
white settlers have replaced the aborigines in the wetter sections
of the same continent.
The Six Faunal Regions
As long ago as 1857, two years before the appearance of
Darwin’s The Origin of Species, an ornithologist named Sclater
published a paper 2 in which he divided the world into six faunal
regions: Ethiopian, Indian, Palearctic, Nearctic, Neotropical, and
Australian. In 1876 Wallace 3 confirmed this division but changed
the name of the “Indian” region to “Oriental”; this change has
persisted in the corresponding literature. Although a century has
passed since Sclater’s work was published, zoologists still divide
the world in this fashion. The faunal regions, which designate the
distribution of the terrestrial and land-locked vertebrates — the
fresh-water fishes, amphibians, reptiles, birds, and mammals —
proved to have been actual divisions during most of the Cenozoic,
or Age of Mammals, except that some of their boundaries shifted
during the glacial and interglacial stages of the Pleistocene epoch.
In earlier times, of course, the surface of the world was divided
differently, but these earlier differences do not concern us in this
book.
Matthew, an influential zoogeographer writing in 1915, 4 indi-
cated that the region of primary evolutionary change was the
2 P. L. Sclater: “On the General Distribution of the Class Aves,” JPLS-Zool.,
Vol. 2 (1857), pp. 130-45.
3 A. R. Wallace: The Geographical Distribution of Animah (London: Mac-
millan & Co.; 1876).
4W. D. Matthew: “Climate and Evolution,” ANYA, Vol. 24 (1915-1939),
pp. 171-318.
THE SIX FAUNAL REGIONS OF
SCLATER AND WALLACE
MAP
5 2 Evolution through Environmental Adaptation
Holarctic, a term combining the Palearctic and Nearctic, and in-
deed those regions were active centers of change during the first
glacial advances of the Pleistocene, when many species of mam-
mals were becoming adapted to cold. However, Darlington now
believes that the tropical regions of the Old World, the Ethiopian
and particularly the Oriental, have been the principal centers of
speciation over a longer period.
The Ethiopian region consists of Africa south of the middle of
the Sahara, which in times of drought acts as a barrier to the move-
ments of many animals, and the southwestern corner of Arabia
south and west of the Arabian desert. But until the end of the
Pleistocene North Africa had an Ethiopian fauna; about ten or
twelve thousand years ago it was invaded by Palearctic mammals,
including Caucasoid men. Madagascar, with an extremely spe-
cialized and archaic fauna, is a special province of its own and
was not inhabited by human beings until about the time of Christ.
South Africa, which lies as far from the equator as South Carolina,
has a Mediterranean climate, but as there is no barrier to separate
it fiom the main part of Africa it has not been isolated enough to
have developed a special fauna of its own. The fresh-water fishes,
amphibia, and reptiles of the Ethiopian region resemble those of
both the Nearctic and the Oriental regions; the birds, as might be
expected, have world-wide relationships, though they are particu-
larly linked to the Oriental region; and the mammals can be di-
vided into certain widely distributed families, some related to the
Oriental region, some purely local, and a few with other con-
nections. However, for the mammals as for the other classes of
land vertebrates, the greatest ties are to be found with the Orien-
tal fauna.
The Oriental region consists of tropical Asia with its fringing
islands, including Ceylon, the Andamans, Sumatra, Java, Borneo,
Formosa, and in certain respects the Philippines. On the east it
encompasses southern China north to Hong Kong, and on the
west it runs a few degrees north of the tropics in northern India.
There is heavy rain forest in much of Indochina, the Malay penin-
sula, Siam, and western Indonesia, and patches of it in the Car-
damon Hills (Kadar country) of southern India and in the Khasi
plateaus of Assam, which is the wettest place in the world.
53
The Six Faunal Regions
Oriental fresh-water fishes form a rich and dominant assem-
blage lacking archaic groups; Oriental amphibia and reptiles are
partly similar to and partly unlike the Ethiopian ones; and
whereas Africa has more species of lizards, the Oriental region is
particularly rich in snakes. Both its birds and mammals are
strongly related to the Ethiopian groups, as for example its ele-
phant, its rhinoceros, and the lion, but the Oriental fauna is less
sealed off than the Ethiopian. It is also related to the Palearctic,
in common with which it has bears and tigers. Both of these are
lacking in the Ethiopian region. This relatively open character is
reflected in the fact that the Oriental region has fewer purely
local (endemic) groups of vertebrates than any other tropical
area. Darlington says: Either it has been a center from which
vertebrates have tended to spread into other regions, or it has
been a main crossroads in dispersal, or both.” 5 Within the Orien-
tal region the fauna can be divided into four regional assem-
blages. The richest and most varied is in the northeastern part,
including southeast China, Indochina, Siam, and Burma; the
poorest is in the principal, drier part of India.
Furthermore, the boundary between the Oriental and Pale-
arctic regions which verges on the richest subarea — the south
Chinese border — is wide open. Nothing except a very gradual
climatic cline stands in the way of free passage northward by
Oriental animals, and vice versa. The width of this frontier is
greatly extended by a series of cool mountain ridges stretching
like fingers from the Chinese highlands southward between the
rivers of southeast Asia. In no other place in the world does an
open border exist between a tropical and a temperate faunal re-
gion. As we shall presently discover, this has been significant for
man as well as for other animals.
On the western side of the Oriental region the mountain bar-
riers are formidable, but there are passes, particularly the Khyber
and Shibar passes, into Palearctic territory, and during certain
warm interglacial periods the Mediterranean and western Europe
had Oriental faunas. The road to the Ethiopian region now runs
along the barren Makran coast of Baluchistan and the connecting
piece of the south Persian coast, then either across a small salt-
5 Darlington: op. cit., p. 436.
54 Evolution through Environmental Adaptation
water gap or around the head of the Persian Gulf, and down into
the Green Mountain of Oman, and finally along the southern
coast of Arabia, where only one undessicated pocket, the Dhofar
region, remains. During the times when the main movements of
animal groups took place between Africa and India, this whole
route must have been much wetter than now and easier to cross.
The Palearctic region includes the nontropical parts of Eurasia
and the Barbary states. Climatically speaking, it ranges from
arctic to Mediterranean conditions, but in all or nearly all of it
there is winter frost, which means that all the animals who live
in it achieved some kind of adaptation to cold. This fauna is
consequently far less rich in species than the Oriental or the
Ethiopian. As the principal flow of animal groups has been from
southeast Asia to China and thence to points north and west, it is
not surprising that the land vertebrate fauna of China is the most
varied of the whole Palearctic region, whereas that of the British
Isles is quite poor. Furthermore, the animals differ more from east
to west than they do from north to south. This is of special interest
to anthropologists, because the same thing is true of human sub-
species. A Norwegian and a Berber resemble each other far more
than either resembles a Chinese, and many a Tibetan could pass
for a Chukchi of northeastern Siberia. The diagonal mountain
barrier running from the Tian Shan range to the Bering Strait,
which partially separates the Caucasoid and Mongoloid realms,
has had its effect on other animals as well.
The Nearctic fauna, which occupies all of North America north
of the tropical part of Mexico, is relatively poor in species, most of
which are derived from the Palearctic, although a few have moved
up from tropical Middle and South America. Greenland is part of
this area, and contains American mammals only, although some
of its birds are European.
South and Central America, the tropical lowland of Mexico,
and Trinidad comprise the Neotropical region. The other islands
of the West Indies have a greatly reduced fauna, which is transi-
tional in a minor way. On the whole, the Neotropical fauna is a
mixture of old forms that developed locally during the Tertiary,
and new ones, including man, which came in from North America
Wallacea
55
during the Pleistocene. A few of its mammals, notably the arma-
dillo and the opossum, have migrated northward.
The Australian faunal region encompasses Australia itself, Tas-
mania, New Guinea, and some of the fringing islands off New
Guinea. That this region has been cut off from the rest of the
world for a very long time is evidenced by the fact that it is
very poor in fresh-water fishes and amphibia, and that its mam-
mals are composed of monotremes (the platypus and echidna)
and of six exclusive families of marsupials. Its closest relationships
are with the almost equally residual fauna of South America, and
it has very little in common with the neighboring Oriental region.
Wallacea
The region between the Oriental and the Australian realms
is named Wallacea. Across it the world’s richest and poorest con-
tinental vertebrate faunas face each other. In i860 Wallace drew
his famous deep-water line between Bali and Lombok, Borneo
and Celebes, and Mindanao and the islands of Sangi and Talaud.
Although Bali and Lombok are only 15 miles apart, the Oriental
fauna is cut off at that point almost as though with a knife. This
line is the western frontier of Wallacea; the eastern is close to the
so-called bird-head, a peninsula of western New Guniea. The
island of Kei is inside Wallacea; and Misol, Waigeo, Batanta, and
Salawati go with New Guinea in the Australian region. A third
line, known as Weber’s, runs down the middle; it is called the line
of faunal balance.
Very few land mammals have crossed Wallace’s Line, and
those that have done so live almost exclusively in the northern
part of Wallacea. Celebes was reached by shrews, tarsiers, ma-
caques, squirrels, four genera of weasels, several kinds of pigs, in-
cluding the endemic Barbirussa deer, and an endemic breed of
cattle known as the anoa. During the Pleistocene Celebes also
harbored a pygmy elephant. Some of these animals reached the
Moluccas, but they did not go south. In the Lesser Sundas, east
of Bali, porcupines, shrews, crab-eating monkeys, pigs, and deer
56 Evolution through Environmental Adaptation
all reach as far as Timor. All these animals may have been intro-
duced by man, who probably brought them along as pets, food,
or both.1’ On the other side, a few Australian marsupials have
penetrated into Wallacea as well. There is a bandicoot on Ceram,
and phalangers live on Halmahera, Timor, and Celebes.
Wallacea is unique in the world as a barrier-filter. It is of great
anthropological significance because it isolated the Australian
aboriginal population virtually unchanged since its arrival from
the southeastern Oriental region during the late Pleistocene.
Even today the population of these small islands is racially inter-
mediate between the more recently arrived Mongoloid peoples of
western Indonesia and the natives of New Guinea. As the ancient
barrier between the erstwhile Sahul and Sunda shelves, it must be
taken into account in any attempt to unravel the complex racial
distributions in southeast Asia and Oceania.
The Faunal Regions and Human Origins and Movements
Man, it is becoming increasingly clear, must have originated
in some form in one of the two realms of tropical fauna in the
Old World, or in an erstwhile extension of one of them into what
is now the Palearctic during a period warmer than the present.
Whereas the exchange of animals between Africa and south Asia
was intermittent and mutual during the Tertiary and Pleistocene,
the Oriental region supplied most of the vertebrate groups to the
Palearctic. One principle of the movements of animals may help
us decide which way the animals moved. When older, less domi-
nant forms are replaced by spreading dominant forms, the domi-
nant ones do not push the others to the peripheries ahead of
them; rather, they overrun them, leaving small, disconnected ref-
ugee pockets in their wake.
The Oriental region contains many such small, marginal popu-
lations of Australoids, Asiatic Negritos, and primitive, food-gath-
ering Caucasoids, which indicates that these races inhabited
that zoogeographic region before its invasion by Mongoloids and
modern kinds of Caucasoids from the north. In the Ethiopian re-
6 Darlington: op. cit., pp. 466-7.
Faunal Regions and Human Origins and Movements 57
gion the distribution of Pygmies follows a similar refugee pattern,
and in East Africa pockets of Bushmen indicate the earlier distri-
bution of Capoids to the north of their historic home. The only
comparable relict population in the Palearctic or Nearctic is that
of the Ainu, and both their antiquity in northern Japan and
their origin are questionable.
Returning to straight zoology, we find that, as many Palearctic
genera originated in south Asia, greater dominance was required
for some of them to move northward through the eastern part of
the Palearctic region and then into Europe and the Americas than
for others to reach the Ethiopian realm, particularly in a period
of greater moisture than the present. Furthermore, as Rensch 7
points out, more evolution has been taking place in the Northern
Hemisphere than in the Southern since the end of Tertiary times,
or in other words, since the beginning of the Pleistocene. As a
great deal of human evolution occurred during the Pleistocene,
the Oriental region, being the more northerly, is a better candi-
date than the Ethiopian as a possible place of dispersal of the re-
mote ancestors of the living races of man.
Certain complications, however, qualify this interpretation. In
the Palearctic region the diagonal mountain barrier that crosses
central and northeastern Asia imposes a bar sinister of cold cli-
mate between the eastern and western halves of this faunal re-
gion. Animals that enter the eastern half from the Oriental region
do not all cross it. During parts of the Pleistocene this barrier was
glaciated. Also, during the Early Pleistocene and the interglacials
of the Middle Pleistocene southern and western Europe were
tropical regions, connected by the Near Eastern land bridge to
both the Oriental and the Ethiopian regions. Asiatic species were
commoner in Europe at these times than African ones, although
both were present.
Therefore, if man did not originate in Europe, which we can
almost take for granted, he could have arrived there from either
southwest Asia or Africa, or from both. It is very unlikely that ini-
tially he came across the mountains of central Asia from China.
At any rate, once Europe had been populated, alternate periods
of glacial and warm or temperate climates gave Europeans more
7 Rensch: op. cit., pp. 275-85.
58 Evolution through Environmental Adaptation
than one chance to adapt themselves to new conditions and, like
other members of their faunas, to reinvade the tropics.
In sum, the rules of zoogeography apply to man as they do to
other animals. They offer us probabilities as to where the genus
Homo evolved and over what paths different groups of men
moved to found regional populations. They also help explain the
dominance of some populations over others.
These rules can be applied in another way as well. Animals of
many species have become adapted morphologically and phys-
iologically to the exigencies of different climates. As members of a
species that inhabits all climates, some of the races of man may
have undergone selection for extremes of climate.
Environmental Adaptation and Early Man
Zoogeography leads by tiny steps into ecology, which
deals with the ways different plant and animal species get along
together in various environments, and ecology in turn carries us
into the study of environmental adaptation. How the polar bear
can sit on a cake of ice without melting it, and how desert rodents
live without water, are fascinating subjects discussed in an exten-
sive literature. The adaptations of living races of men to climatic
extremes have also been studied, to a lesser extent. But for present
purposes, since we are concerned only with the history of fossil
human races, only two aspects of this subject need be explored
here.
( 1 ) We need to know whether the fragmentary remains of our
fossil ancestors contain any telltale indications of adaptation to
climatic extremes, in order to determine whether such adapta-
tions aie chaiacteristic of subspecies, and to keep ourselves from
confusing them with general, evolutionary characters.
(2) We need to determine what extremes of climate our an-
cestors could have tolerated with a minimum of cultural equip-
ment and we can do this by studying and comparing the physi-
ology of living primitive peoples.
The results of both these investigations may help us determine
how old the existing subspecies are, and whither and whence they
could have migrated during the Pleistocene.
The Rules of Bergmann and Allen 59
Simply by observing the geographical distribution of living
peoples, we can see that no single subspecies is limited to a single
climate. Caucasoids live all the way from Norway to India. The
aborigines of Tasmania, who were spiral-haired Australoids, went
about nearly naked in a climate as cold as England’s. Mongoloids
may be found from the Arctic to the wet tropics, and both the
Australian aborigines and the South African Bushmen, whose
ranges are more limited, live through broiling heat and freezing
weather at different seasons, with a minimum of cultural assist-
ance. Given time, a population derived from any human sub-
species could probably adjust itself to most local conditions.
Adaptations to climate, therefore, could occur independently in
more than one subspecies and need not be interpreted as evidence
of genetic relationship.
In only three kinds of environment are land mammals rigor-
ously selected for their abilities to resist stress: arctic and other
cold areas, deserts, and high mountains. The first entails heat
regulation; the second both heat regulation and water conserva-
tion; and the third oxygen consumption, particularly oxygen
transfer from the mother to the fetus. Physiologists have done a
great deal of work to explain how the caribou can live in the snow,
why the camel can go for days in the summer heat without drink-
ing, and how the llama can bear its young in the thin air of the
Andean plateau.
The Rules of Bergmann and Allen
These adaptations found among living mammals in-
volve fur, skin, blood vessels, interstitial fluids, and blood cor-
puscles, i.e., soft parts, which ordinarily disappear after death. If
geologically ancient human beings also had such adaptations,
very few details of these can be detected among the bones at our
disposal. Yet certain uniformities which reflect relationships be-
tween the bodies of animals and climate may show in the skeleton
as a whole. These are the old, nineteenth-century rules. Berg-
mann s rule,8 for example, states that in a given species the warm-
8 Carl Bergmann: Uber die Verhaltnisse der Warmeokonomie der Thiere zu
ihrer Grosse,” Gottinger Studien, No. 8 (1848).
60 Evolution through Environmental Adaptation
blooded animals which live in cold places tend to have greater
body bulk than those which live in hot regions. Allen’s rule 9
further states that in a given species animals living in cold areas
tend to have shorter extremities than those in warm climates.
This does not mean that in cold places animals’ ears, legs, or tails
become too short to function, only that within functional limits
they will become shorter than they might have been had cold
stress been absent.
Both these rules concern the physics of heat loss from a warm
body to a usually cooler surrounding atmosphere. As a body has
three dimensions and its surface only two, the bigger the body,
all else being equal, the less the heat loss per unit of volume
(Bergmann). Furthermore, the nearer the body comes to being a
perfect sphere, the smaller is its surface area per unit of volume
(Allen). Each species of animal usually has its own system of
conserving and losing heat, so that these rules cannot be used in
interspecific comparisons.
In recent years these venerable rules have been criticized by
physiologists, some of whom were unaware that the rules apply
only to single species; and they have been defended by taxono-
mists and physical anthropologists.1 As they represent results
rather than processes, they are naturally less useful in studying
adaptation than physiological experiments are, but they can be
applied to much larger population samples than physiologists can
test. As a supplement to physiological experiments, they can be
applied to living men, particularly to old, long-established food-
gathering populations.
Peoples who live in cold regions are generally heavier than the
9 J. A. Allen: “The Influence of Physical Conditions in the Genesis of Species,”
RR, Vol. 1 (1877), pp. 108-40. (Reprinted in ARSI for 1905 [1906], pp.
375-402. )
1 For this controversy see:
P. F. Scholander: “Evolution of Climatic Adaptation in Homeotherms,” Evo-
lution, Vol. 9, No. 1 ( 1955), pp. 15-26.
Scholander: “Climatic Rules,” Evolution, Vol. 10, No. 3 (1956), pp. 339-40.
Mayr: “Geographical Character Gradients and Climatic Adaptation,” Evolu-
tion, Vol. 10, No. 3 ( 1956) pp. 105-8.
M. T. Newman: “Adaptation of Man to Cold Climates,” Evolution, Vol. 10,
No. 3 (1956), pp. 101-5.
C. G. Wilber: “Physiological Regulations and the Origins of Human Types,”
HB, Vol. 29, No. 4 (1957), pp. 329-36.
The Rules of Bergmann and Allen 61
inhabitants of the tropics, and the ratio of trunk length to leg
length is greater in the peoples who dwell in cold areas, who
weigh more per unit of stature.2 As expected, these regional dif-
ferences are found in all subspecies that encompass wide ranges of
climate.
For our present purpose of detecting climatic adaptation in
fossil men these rules are rarely useful, with a few exceptions.
Several nearly complete skeletons of European Neanderthals have
bones so short and heavy that their body weights must have been
great per unit of stature, as with living peoples of the Arctic.
With these exceptions, we rarely have enough bones from a single
individual to calculate both stature and relative trunk height; in-
deed, stature is usually calculated from the limb bones alone,
which defeats our purpose. Also, there is no formula for calculat-
ing body weight from the skeleton.
Now and then we find the cervical vertebrae, which tell us
whether necks were long or short. This is useful because peoples
in cold climates tend to have short necks. We can also estimate the
amount of warm arterial blood that flows into the cheeks
through the infraorbital foramen (a hole in the zygomatic bone
just under the eye socket) by the diameter of that opening. A
strong flow of blood through that hole helps keep the cheeks of
the Greenland Eskimo warm.3 Similarly, the size of the mental
foramen (mental means chin in this case), a comparable hole in
the lower jaw, affects the amount of warm blood that reaches the
chin.
The shape of the foot is also significant, for people who go bare-
foot in cold water or snow tend to have short broad feet with
short toes. In a few sites whole feet of fossil men have been re-
covered; in others footprints have been found.
Among living peoples who dwell near or above the Arctic
Circle, whether they are Caucasoid or Mongoloid, there is a
tendency for the tympanic plate, a bony structure below the ear
2 D. F. Roberts: “Body Weight, Race, and Climate,” AJPA, Vol. 11, No. 4
(i953), pp. 553-8.
Newman: The Application of Ecological Rules to the Racial Anthropology
of the Aboriginal New World,” AA, Vol. 55, No. 3 (1953), pp. 311-27.
3 W. S. Laughlin and J. B. Jprgensen: “Isolate Variation in Greenlandic Eskimo
Crania,” ActG, Vol. 6 (1956), pp. 3-12.
62 Evolution through Environmental Adaptation
hole, to become thickened — why we do not know. This thickening
has also been observed among the Moriori, the original Poly-
nesian inhabitants of the Chatham Islands, who lived in a cool
climate.
Nose Form and Climate
A further adaptation concerns the nose. In places where
the air is dry the nasal aperture tends to be narrow; where it is
damp, the openings may be broader. This adaptation involves the
function of the nasal passages in moistening inhaled air. Noses
also tend to be narrower in cold than in hot climates, because of
the heat exchange between the lungs and the inhaled air, but the
protection of the lungs from frost is not as critical as the humidi-
fying function.4
Ridges and surface irregularities on the skull and mandible in-
dicate how much and how hard a prehistoric individual chewed.
This can also be determined, in mature specimens, from the
amount of tooth wear. In the earliest fossil hominid remains, par-
ticularly those from periods and places without fire, powerful jaws
and large, heavily worn teeth reflect a coarse diet without clean-
ing, cooking, or other effete ways of demineralizing or softening
food. Later on, in advanced prehistoric populations living in cold
places, jaw muscles (as indicated by the effects they left on bone)
again became massive and teeth excessively worn. Like the Eski-
mo, these people used their teeth in preparing skins for clothing.
Physiological Adaptation to Cold
These imperishable details of skull morphology tell us much
less about adaptation to climate than the soft parts would have
done had they been preserved, as those of mammoths were. Cli-
matic adaptation is physiological, and the physiology of heat and
cold adaptation is mostly a matter of oxygen consumption, blood
flow, and details of muscles, fat, skin, and nervous tissue. Because
4 A- Thomson and D. Buxton: “Man’s Nasal Index in Relation to Certain
Climatic Conditions, JRAI, Vol. 53 (1923), pp. 53—92.
J. S. Weiner: “Nose Shape and Climate,” A]? A, Vol. 12, No. 4 ( 1954), pp. 1-4.
Physiological Adaptation to Cold 63
differences in physiology are racial, and racial differences are as
old as Homo sapiens, we may venture to project physiological dif-
ferences in living races backward into the time of fossil men.
Many such differences, long suspected, have recently been estab-
lished.
During the 1940’s the global nature of modern warfare stimu-
lated the interest of several nations in man’s ability to live in all
climates, particularly the arctic. It soon became clear to some
researchers that living races differ in their tolerance of heat and
cold. Although much work remains to be done, at least seventeen
experimental studies published between 1950 and i960 reported
tests of this nature on all five subspecies of Homo sapiens.5 6
These tests have shown that Mongoloids are adapted to sleep-
ing and working in the cold as a result of one kind of physiological
adaptation; that Australoids and one group of Caucasoids, the
Lapps, are cold-adapted in an entirely different way; that Ne-
groes are both adapted to wet heat and sensitive to cold; and
that most European Caucasoids and all Bushmen studied lack
special adaptations to either heat or cold.
In the Arctic, fur keeps the bodies of most mammals warm.
The same furs, tailored into clothing, keep people warm out of
doors. But despite the use of warm clothing, blankets, and camp-
fires, Alaskan Indians sleep under conditions of moderate cold
while camping out on their trapping routes in the winter. Then-
bodies, however, compensate for the incurred heat loss by an in-
creased basal metabolism. By burning extra oxygen and calories
they are able to sleep without discomfort at temperatures that
keep white men tossing and waking.0 This physiological capacity,
which is inherited, is not a seasonal phenomenon. It keeps them
warm, with little cover, on chilly summer nights as well as in
winter.7
5 Europeans were used as controls in all these experiments except that given in
footnote 3, page 65, the Japanese tests. Thus Caucasoid Europeans, other than
Lapps, who were tested separately, constitute the norms.
6 L. Irving, K. L. Anderson, A. Bolstad, R. Eisner, J. A. Hildes, Y. Lpyning, J. D.
Nelms, L. J. Peyton, and R. D. Whaley: “Metabolism and Temperature of Arctic
Indian Men During a Cold Night,” JAP, Vol. 15, No. 4 (i960), pp. 635-44.
R. W. Eisner, K. L. Anderson, and L. Hermanssen: “Thermal and Metabolic
Responses of Arctic Indians to Moderate Cold Exposure at the End of Winter,”
JAP, Vol. 15, No. 4 (i960), pp. 659-66.
64 Evolution through Environmental Adaptation
A far more spectacular and much better known example of cold
adaptation is that of the Canoe Indians of Tierra del Fuego and
adjacent South American shores and islands. In 1959 Hammel,
Scholander, and others, including myself, went to the islands and
glaciers of the southern Chilean archipelago to study the cold
adaptation of the Alakaluf,8 the only one of the four original
Fuegian tribes still numerous enough and unmixed enough to war-
rant investigation.
When first discovered by Magellan, these Indians were going
about in canoes in freezing weather with no clothing except an
occasional sea-otter skin cape, and with their bodies smeared with
sea-mammal fat and ocher. At night they usually slept in small,
domed huts covered with skins and heated by fires of Nothofagus,
an evergreen tree closely related to the beech. This wood throws
oflf great heat and burns nearly all night.
Except for the early morning hours, these Indians were as warm
indoors as we are. Out of doors they exposed themselves un-
clothed to heavy winds and pelting sleet and snow. Furthermore,
they walked and swam in the icy water, and dived for shellfish.
The work of Hammel and his associates shows that the Fuegians,
taking the Alakaluf as an example, were able to survive freezing
temperatures without clothing by burning off a large quantity
of calories, much more than the Alaskan Indians needed to keep
warm at night. The Alakaluf live mostly on shellfish and the flesh
of sea mammals, and they eat heartily. Their basal metabolism is
160 per cent higher than the norm for whites of the same weight
and stature.
Returning for a moment to arctic mammals, and also to arctic
birds, we observe that no matter how warm their fur and down
keep their bodies, certain extremities, like seals’ flippers, caribou’s
lower legs, and birds’ beaks, remain relatively unprotected. In
some species, as for example the fur seal with its exposed flippers,
warmth is provided to these extremities by a massive flow of
arterial blood close below the surface. This flow of blood burns
up many calories, thus enabling the seal to swim in comfort.
On anatomical evidence alone, we have already inferred that
s H. T. Hammel: Thermal and Metabolic Responses of the Alacaluf Indians to
Moderate Cold Exposure, WADD Technical Report 60-633, December i960.
Physiological Adaptation to Cold 65
the cheeks of the Greenland Eskimo receive an extra flow of blood
which keeps them warm, but as far as I know this has not yet been
tested physiologically. The Eskimo’s hands, however, have been
tested for the same phenomenon, and they show an increased
flow of blood when held in cold water.9
The hands of Alaskan Indians respond in the same fashion,
producing twice as much blood flow as those of white men tested
under the same conditions.1 The same response was obtained
from the hands of Alakaluf women,2 who collect shellfish by hand
in cold water. In Manchuria four groups of Mongoloids were
tested by the Japanese for this same phenomenon,3 and a grada-
tion, or cline, was found which corresponds to the climates of the
regions inhabited by the peoples studied. The Orochons, a no-
madic, reindeer breeding and hunting tribe of northern Man-
churia, had the most adaptation; the Mongols and north Chinese
came next (the two were the same); and the Japanese had the
least response.
Similar tests performed on the hands of Lapp reindeer herders,
who have been living since prehistoric times under the same con-
ditions as the Orochons, showed no cold adaptation in the hands.4
White Norwegian fishermen living above the Arctic Circle, men
whose hands are constantly in cold water, came out the same as
the Lapps, and as the white men in the control group, who were
mostly scientists.5
The experiments reported above indicate that cold adaptation
9 G. M. Brown and J. Page: “The Effect of Chronic Exposure to Cold on
Temperature and Blood Flow of the Hand,” JAP, Vol. 5, No. 5 ( 1953), pp. 221-7.
1 Eisner, Nelms, and Irving: “Circulation of Heat to the Hands of Arctic In-
dians,” JAP, Vol. 15, No. 4, pp. 662-6.
2 H. T. Hammel: “Thermal and Metabolic Responses. . . .”
3 H. Yoshimura and T. Iida: “Studies on the Reactivity of Skin Vessels to
Extreme Cold. Part II: Factors Governing the Individual Difference of the Reac-
tivity, or the Resistance Against Frostbite,” JJP, Vol. 1 (1950-51), pp. 177-85.
4 J- Krog, B. Folkow, R. H. Fox, and Andersen: “Hand Circulation in the Cold
of Lapps and North Norwegian Fisherman,” JAP, Vol. 15, No. 4 (i960), pp.
654-8.
B. Hellstrom and Andersen: “Heat Output in the Cold from Hands of Arctic
Fishermen,” JAP, Vol. 15, No. 5 (ig6o), pp. 771-5.
5 Ibid.
Andersen, Lpyning, Nelms, D. Wilson, Fox, and A. Bolstad: “Metabolic and
Thermal Response to a Moderate Cold Exposure in Nomadic Lapps,” JAP, Vol.
15, No. (i960), pp. 649-53.
66 Evolution through Environmental Adaptation
through increased basal metabolism and increased peripheral
blood flow is confined to the Mongoloid subspecies, at least as
far as we know. They also indicate that the Lapps are Caucasoids,
as most physical anthropologists now believe, and not Mongoloid,
as was frequently stated in the past by writers who had not seen
them.
The second kind of cold adaptation requires no increase in
caloric expenditure or in peripheral blood flow. It involves instead
an insulation in depth of the body core; the limbs and the sur-
faces of the trunk serve to insulate the more vulnerable internal
organs. This effect is found in domestic swine reared in Alaska,
and in hair seals, which have no more fur than the swine do.
It is also characteristic of the legs of caribou and of arctic birds.
Like the Mongoloid adaptation, this type involves both the body
as a whole and the extremities.
In man, cold adaptation through insulation was first observed
in Australia, among the aborigines. In west-central Australia the
members of the Pitjendjera tribe live naked in the desert. During
the day the air is hot, but at night the temperature can go down
to freezing or a little lower. Ordinarily the aborigines sleep naked
on the ground between rows of small, smudgelike fires, but when
the wind is blowing the fires are useless. Scholander, Hammel, and
others found that, while sleeping in light sleeping bags without
fires at 32 ° F, the Pitjendjera men maintain an almost normal in-
ternal body temperature, as shown by rectal readings, whereas
their limbs become chilled. The temperature of their feet read as
low as 540 to 590 F.
In the morning these men get up and stamp around, and by
the time the sun is up they are as fit as ever. White volunteers
who took the same tests lost internal body heat before morning,
because the surfaces of their arms and legs threw it off into the
atmosphere. The aborigines slept comfortably, but their Cauca-
soid counterparts spent a miserable night.6 Later on, these experi-
ments were repeated in midsummer at Darwin, North Australia,
on other aborigines from several different tribes. Cold condi-
6 P. F. Scholander, Hammel, J. S. Hart, D. H. LeMessurier, and J. Steen:
“Cold Adaptation in Australian Aborigines,” JAP, Vol. 13, No. 2 (1958), pp.
211-18.
Physiological Adaptation to Cold 67
tions were created by having them sleep in a refrigerated meat
van. The physiological response was the same as that of the first
group, tested in winter, thereby confirming the fact that the cold
adaptation of the Australian aborigines is not seasonal but per-
manent, and apparently both genetic and anatomical.7
In human beings each of the principal arteries of the arm and
lower leg — brachial, radial, ulnar, tibial, and peroneal — is ac-
companied, as a rule in the same sheath, by a pair of companion
veins called venae comites. At various places, particularly near
elbows and other joints, neighboring arteries are connected by
short blood vessels, so that under certain circumstances one can
replace the other and an exchange of blood can take place. The
networks formed by such connections are called anastomoses.
The economy of engineering that placed the arteries and their
pairs of veins together, and the emergency arrangement of con-
necting arteries at anastomoses, have also provided a mechanism
by which under certain circumstances heat can be transferred
between the two kinds of blood vessels.
Among the Pitjendjera apparently such a transfer is made dur-
ing sleep. The outgoing arterial blood warms the incoming venous
blood, so that the hands and feet are cool and heat is saved. In a
desert where food is scarce, heat conservation is important for
survival. Why the whites tested in these experiments failed to
transfer heat from arteries to veins in the same way is not known,
but without doubt a program of comparative dissection could
help determine the answer. Arteries are notoriously variable, and
racial differences in their branching patterns have been estab-
lished between Europeans and Japanese. For other populations,
available data are inadequate.8
Surprisingly enough, the Australoid type of cold adaptation
through insulation has been found in only one other population
so far tested, the nomadic Lapps. This evidence, when added to
their failure to respond to the cold-water hand test, places the
Lapps far from the Mongoloid subspecies. Also, the settled village
7 Hammel, Eisner, D. H. LeMessurier, Andersen, and F. A. Milan: “Thermal
and Metabolic Responses of the Australian Aborigine Exposed to Moderate Cold
in Summer,” JAP, Vol. 14, No. 4 ( 1959), pp. 605-15.
8E. Loth: L’ Anthropolo gie des Parties Molles (Warsaw and Paris: Masson et
Cie; 1931), PP- 348-82.
68 Evolution through Environmental Adaptation
Lapps, who are more mixed with Finns and Norwegians, show the
insulative cold adaptation less than do the reindeer herders, who
are less mixed. One is tempted to suspect that this type of cold
adaptation was prevalent in Europe during the latter part of the
Wiirm glacial epoch.
Returning to the Southern Hemisphere, where physiologists
have found the world’s most striking examples of cold adaptation
— among the Fuegians and Australian aborigines — we approach
the Bushmen of the Kalahari Desert with some hope. The Bush-
men, who are also primitive hunters and gatherers, are faced with
the same alternate stresses of heat and cold that confront the
Australian aborigines. These hopes have not been realized, how-
ever. Three separate expeditions 9 have failed to find any differ-
ences, in basal metabolism or in any other physiological attribute,
between the Bushmen and the whites used as controls. This evi-
dence suggests what has been suspected on other grounds, that
the Bushmen have not lived in the desert very long. It also con-
firms my belief that the Bushmen and the Negroes, although they
share a continent, are not closely related.
Heat Adaptation
S o far, only the Negroes have been shown to possess heat
adaptation. American Negores can tolerate moist heat better than
American whites of the same age and economic background.1 But
as far as I know this difference has not yet been demonstrated
in Africa.2 American Negroes are unable to tolerate cold as well as
9 C. H. Wyndham and J. F. Morrisson: “Heat Regulation of MaSarwa” (Bush-
men), Nature, Vol. 178, No. 4538 ( 1956), pp. 869-70.
Wyndham and Morrisson: “Adjustment to Cold of Bushmen in the Kalahari
Desert,” JAP, Vol. 13, No. 2 (1958), pp. 219-25.
J. S. Ward, G. A. C. Bredell, and H. G. Wenzel: “Responses of Bushmen and
Europeans on Exposure to Winter Night Temperatures in the Kalahari,” JAP,
Vol. 15, No. 4 (i960), pp. 667-70.
1 P. T. Baker: “Racial Differences in Heat Tolerance,” AJPA, Vol. 16 (1958),
pp. 287-305.
T. Adams and B. G. Covino: “Racial Variations to a Standardized Cold Stress,”
JAP, Vol. 12, No. 1 (1957), pp. 9-12.
2 A study conducted in West Africa by N. A. Barnicot yielded negative re-
sults, possibly because he apparently failed to allow for differences in height.
The Significance of Adaptation to Heat and Cold 69
American whites; this is true even when the individuals of both
races who are tested have the same amount of subcutaneous fat.3
This final observation indicates that the difference in thermal
adaptation between Negroes and European Caucasoids is not due
to insulation alone. Probably a whole complex of physiological
processes is involved, particularly those concerned with the depo-
sition of melanin in the skin by the action of three hormones.4
The Significance of Adaptation to Heat and Cold
Several conclusions can be drawn from this review of human
adaptation to heat and cold. One is that the subspecies of man as
defined in Chapter 1 tend to sort themselves out on this basis. The
Mongoloids are the most distinctive in thermal adaptation as in so
many other features, and the Negroes stand at the opposite ex-
treme.
A second is that because these adaptations are both genetic and
linked to climate they may have been acquired by the several
subspecies of Homo erectns at the time of their dispersal into
different environmental regions.
A third conclusion is suggested by the Alakaluf study. It indi-
cates that ill-clad human beings carrying fire and the crudest of
tools (the Alakaluf cutting tool was a quahaug shell) could have
entered North America over the Bering Strait at any time when
the sea level was low enough to permit passage. At such times,
with the flow of arctic water cut off and the Japanese current
swinging along the southern shoreline, the climate could have
been no colder than it is in modern Tierra del Fuego.
The two kinds of cold adaptation recently discovered allow
weight, and bodily components between the Negroes and Europeans tested. N. A.
Bamicot: “Climatic Factors in the Evolution of Human Populations,” CSHS, Vol.
24 (i959), PP- 115-29.
3 Baker: “American Negro-White Differences in Thermal Insulative Aspects of
Body Fat,” HB, Vol. 31 (1958), pp. 287-305.
4 Melanin is deposited by the combined action of one hormone from the pineal
gland and two from the pituitary. The melanocytes in which the pigment is
formed have their embryonic origin in nerve cells. Thus, skin pigment is basically
a neuroendocrinological product. A. B. Lerner: “Hormones and Skin Color,” SA,
Vol. 205, No. 1 (1961), pp. 98-108.
7° Evolution through Environmental Adaptation
human beings to live at temperatures near the freezing point with
little or no environmental protection when out of doors, but they
would not allow anyone, however well adapted genetically, to
hunt out of doors in the winter temperatures found today in Lap-
land and Greenland without a combination of good clothing and
good housing, both made with good tools by skilled hands di-
rected by a fully evolved modern brain. As far as the fossil record
tells us, only Homo sapiens has ever lived in such climates.
Adaptation to Altitude
And as far as we know only Homo sapiens has ever lived at
altitudes of over 10,000 feet. Only two plateaus of this height
which are large enough to be human breeding grounds exist.
They are Tibet and the Andean altiplano. Both are inhabited by
Mongoloids. Careful physiological and anthropometric work has
shown that the Andean Indians have large chests, large lungs,
large hearts, and blood that contains a high ratio of red corpuscles.
Although each red corpuscle carries less oxygen than it would at
sea level, the total amount of oxygen borne by the blood far ex-
ceeds that supplied by the arteries of outsiders who have moved
into the highlands. Such outsiders may survive, but they have
difficulty reproducing because the mother cannot transfer enough
oxygen to her embryo to ensure its live birth.5 That is one reason
why the highlands of Ecuador, Peru, and Bolivia are still Indian
country four and a half centuries after Pizarro. As far as I have
been able to determine, the adaptation of Tibetans to high alti-
tudes has not yet been studied.
On the opposite extreme, Negroes, whose blood carries the sick-
ling trait polymorphically ( Ss ) and bears with it even less oxygen
than that of Caucasoids, may be seen along the Andean coast but
not on the plateau. In the Himalayan region the clinal zone be-
tween Mongoloids and all others is extremely steep, and in some
places it is only a few miles wide.
The fact that adaptations favoring or counteracting excesses of
oxygen in the blood stream cannot be demonstrated in fossil man
5 Newman: “Man and the Heights,” Nil, Vol. 67, No. 1 (1958), pp. 9-19.
Adaptation to Altitude yi
does not mean that they did not exist, because such adaptations
are found only in perishable fluids and tissues.
In this chapter we have surveyed the principles of geography
as they may be applied to the distribution of animals and the de-
velopment of species and subspecies. We have situated the sub-
species of man in their ancient homes, and examined the evidence
for climatic adaptation in fossil and living men. We have found
that human subspecies differ considerably in climatic adaptation,
which has played a part in the ability of human beings to invade
and inhabit regions too cold or too dry for other primates. The
historic distribution of races, in fact, may partly be explained on
the basis of these adaptations.
But we have found no extreme forms of adaptation comparable
to those of desert rodents that live without drinking water, or of
polar bears that sleep naked on ice floes. The principal adapta-
tions of human beings to climate are technological. Skill at tech-
nology’ and particularly the inventive genius that makes technical
advances possible, requires the possession of a top-grade brain,
which our ancestors began to acquire long ago, and which is still
useful in an increasingly technological society.
« &
EC 3 K
EVOLUTION THROUGH SOCIAL
ADAPTATION
Leadership , Communication, and Brain Growth
top-grade brain is needed not only to master, by
technical means, cold, drought, and other environmental difficul-
ties beyond the physiological capacities of the human body, but
also to manage human relations skillfully. Natural selection in
favor of this second kind of skill has been a prime factor in hu-
man phyletic evolution — the rise of a more intelligent species
from one that is more primitive intellectually. In this chapter I
shall try to show how this kind of natural selection may have
operated.
I am particularly concerned with the surviving societies of
primitive hunters and gatherers because they serve, to a certain
extent, as a window into the distant past, but more advanced
systems should not be neglected since all societies are governed
by the same natural rules.
In all the historic societies whose structural details are well
known, the greatest tangible rewards have rarely gone to the
geniuses of technology or to outstandingly skilled craftsmen, how-
ever important their work has been for the preservation of human
life and to social evolution. The men who have reaped the highest
rewards are the geniuses, artists, and skilled craftsmen whose ma-
terial is not clay, flint, or metal, but other people. They are the
“operators,” the artificers of human relations. The leader who can
keep the peace among his followers, organize his men for war,
regulate the distribution of food and other wealth in such a way
73
Leadership, Communication, and Brain Growth
that everyone will be taken care of, particularly himself — such a
man is well paid. He lives in the finest structure, be it hut or
palace, eats the best food, and in many societies has the most
women. Whatever genes he has that others lack have a better
than average chance to multiply in the local pool.
Also well rewarded in esteem, if not in material goods, is the
priest, shaman, or medicine man whose artistry allays fears and
eases people individually and as groups over the emotional
hurdles of crisis and trouble. In many societies his personality is
an odd one. As he ministers to both sexes, he is sometimes celibate.
What makes him an artist does not necessarily give him more
women than the others; a society in which everyone is a shaman
would soon fall to pieces. A few of his special genes in the pool
will go a long way. Like popes, he can pass on his heritage
through nephews.
Under the umbrella of law and order, ritual sanction, and emo-
tional security that both chief and shaman spread, the craftsman
can do his work, and every man can get food for his family. As
there must be leaders, there must also be followers — men and
women who can live together under guidance without disruptive
quarreling. During the long stretch of human evolutionary history
the sizes and complexities of groups have grown, and the ability
of group members to live together peacefully, while presenting a
united front against outsiders, has been of great importance for
survival. In many structurally simple societies the troublemaker
is killed one dark night by his fellows, or driven away, and so
the genes which may have contributed to his antisocial behavior
are thus, in a sense, fished out of the pool. Social adaptation,
which is the capacity for living together in groups, has been as
influential in human evolution, if not more so, as environmental
adaptation through technology. But the relative importance of
these two facets of adaptation is hard to evaluate as they are parts
of a single picture.
Both these categories of adaptation depend primarily on an
ancient revolution in communication made possible by the inven-
tion of speech. Like tool-making and the use of fire, speech, we
know, was a human invention. It must be learned, not quickly
like some of the semi-instinctive habit patterns of other mammals
74 Evolution through Social Adaptation
but slowly and with great effort, and it requires the co-ordination,
within the brain, of several different organs that are not used in
concert by any other primate. If speech did not have to be
learned, the peoples of the world would not speak hundreds of
different languages; they would all make the same noises, like
sea gulls.1
Before speech could be invented, the ancestral primate or-
ganism had to undergo certain anatomical changes.2 These involve
the following organs of speech (and, of course, their nerves):
the diaphragm, which expels the air from the lungs; the larynx,
which contains the so-called vocal chords and their controlling
muscles; the pharynx, which is essentially the valve that opens
and shuts the intersection of the air and food passages of the
throat, both below and above the meeting point; and the muscles
that control the movements of the jaws, lips, tongue, and soft
palate.
The principal change was in the pharynx. In primates that walk
on all fours, the air tube is continuous from pharynx to nasal
passages except when the animal is swallowing or crying out; it
takes effort to expel breath through the mouth. In man the valve
of the pharynx is habitually open, and breath will come out of
the mouth whenever the lips are open and the lungs are exhaling,
unless an effort is made to block its passage with the tongue.
The cause of this change was, apparently, the assumption of
the erect posture by our ancestors. DuBrul has shown by a series
of dissections of the heads and necks of tree shrews, lemurs,
tarsiers, Old World monkeys, and apes that the opening of the
pharynx in man was only the last step in a series of changes
caused by an increasing postural shift from the horizontal to the
vertical plane. In the most primitive primates the air passages
form almost a straight line from lungs to lips. In man they are
1 For a thorough discussion of the origin of speech and its role in cultural
evolution, see:
A. I. Hallowell: “Self, Society, and Culture,” in S. Tax: Evolution After Dar-
win (University of Chicago Press; i960), pp. 309—71.
C. F. Hockett: “The Origin of Speech,” SA, Vol. 203, No. 3 ( i960), pp. 88-96.
2E. L. DuBrul: Evolution of the Speech Apparatus (Springfield, 111.: Charles C
Thomas; 1958); and “Structural Evidence in the Brain for a Theory of the Evo-
lution of Behavior,” PBM, Vol. 1, No. 4 (i960), pp. 40-57.
Leadership, Communication, and Brain Growth 75
bent, in the pharyngeal section, into a 45 0 angle. It was this
bending that opened the valve.
Once the pharynx was open, air was free to move between
larynx and lips, whether the flap of the soft palate had closed off
the nasal passages or left them open. Now it was possible to utter
a wide variety of sounds, the formation of which depended on a
combination of many factors: the degree of tension of the vocal
cords, which could either be tightened so as to vibrate and thus
Dots = breathing tube
Solid = feeding tube
Fig. 2 The Speech Organs of Pri-
mates. A. Lemur rufifrons. The soft
palate overlaps the epiglottis, and the
corniculate cartilage of the pharynx is
hooked to hold its grip over the rear
rim of the palatal additus. Air pas-
sages are normally open and food
passages closed except in swallowing.
B. Homo sapiens. The larynx has slid
far down the neck. Both the front and
rear valves are normally open, permit-
ting free air to flow into the oral cav-
ity, while the back flap of the soft
palate can close off the nasal passages
in speaking. ( Drawings after DuBrul,
1958.)
emit voiced sounds, or left slack so as to permit the formation of
unvoiced sounds which, if continuous, became whispering; the
opening and closing of the nasal passages, which produce nasal
sounds if left open; the positions taken by the tongue and lips;
and the sequences of all these elements in the formation of
words. The number of possible sounds is nearly infinite, but the
7 6 Evolution through Social Adaptation
number used in any one language is limited by the number that
can be easily recognized.
To be understood, language must be heard, both by the speaker
and by the person addressed. The vocal vibrations of speech pass
into the outer environment and return to the brain through the
ears. If successful communication is achieved, they also hit the
eardrums of a second person, whose answers strike the eardrums
of the originator of the conversation.
Speech requires the neural co-ordination, in the brain stem and
cortex, of many organs and sets of muscles, all of which, being
located near the brain, enter it independently, as do the auditory
nerves, rather than through the spinal cord. Their co-ordination
in the brain was different neurologically from that of the hands
and eyes needed for tool-making. Also, it was acquired later than
the hand-eye combination that brachiation (swinging from limb
to limb) called for: an ape has to see where he is going, in order
to place his hand, or he will fall.3
Therefore, speech was probably invented after tool-making.
Tools made hunting possible, and the social requirements of a
group of hunters made speech necessary. Speech is also a pre-
requisite to thinking, because we think in words. He who thinks
can plan ahead, and he who plans ahead can learn to deal with
other human beings.
During the course of human evolution, in different parts of
the world, the brains of successive fossil men grew larger as time
went on, until the present brain sizes, typical of the living races
of man, were reached. Undoubtedly, talking and thinking influ-
enced these increases, which occurred as more and more had
to be learned. Evolutionary increases in brain size have not been
confined to man. The fossil record shows comparable changes in
many other kinds of animals. What is unusual about man is not
that his brain grew, but that it grew as much as it did.4 By and
3 The other primates lack the extensive pharyngeal plexus needed for speech
which is found in man. J. M. Sprague: “The Innervation of the Pharynx in the
Rhesus Monkey and the Formation of the Pharyngeal Plexus in Primates,” AR,
Vol. 9°, No. 3 (1944), PP- 197-208.
4 For the problem of brain size vs. body size in animals, see:
B. Rensch: “The Relation Between the Evolution of Central Nervous Func-
Leadership, Communication, and Brain Growth 77
large, in response to the needs of communication, the growth of
the human brain may be considered primarily a social adapta-
tion and, in addition, an example of evolution through succession.
This increase in brain size probably started with the erect
posture. In any evolutionary line of mammals any entirely new
kind of locomotion must be learned. Baby seals, for example, must
be taught to swim, and baby birds must be pushed out of their
nests before they will fly. Each of us, as a baby, must be taught to
walk, or we would go on all fours. Learning a new method
of locomotion fosters, and indeed requires, a concomitant increase
in intelligence and, by the same token, in brain size. An animal
bright enough to learn to walk erect might also be bright enough
to begin making tools, and so on to hunting and speech.
But brain growth has disadvantages that had to be outweighed
by the greater advantages of an increasing intelligence. In the
fossil record of our zoological family, brain size increased only
gradually; our brain is an expensive organ that grew as man be-
came increasingly able to support it. The brain requires a large
skull that must be carried about by the bones, tendons, and
muscles of the neck, trunk, and legs. Being very sensitive to
changes in temperature, it must be kept warm in cold weather
and cool in hot weather. Only the visceral organs, which are much
better insulated by the body mass, require such a narrow thermal
range. As the brain lies close to the surface of the head, its large
size taxes the body’s capacity for maintaining thermal equilib-
rium.
It is also a gluttonous organ, requiring an even blood flow
ranging from about 765 cc. a minute when at rest to about 1300 cc.
a minute when hard at work. At rest it monopolizes about 12 per
cent of the body’s blood supply, although it comprises only about
tions and the Body Size of Animals,” in J. Huxley, ed.: Evolution as a Process
(London: Allen & Unwin; 1954), pp. 181-200.
H. J. Jerison: “Brain to Body Size Ratios and the Evolution of Intelligence,”
Science, Vol. 121, No. 3144 ( 1955), pp. 447-9.
Rensch: “Trends Towards Progress of Brains and Sense Organs,” CSHS,
Vol. 24 ( 1959), pp. 291-303.
For the functioning of the brain, particularly in speech, see W. Penfield and
L. Roberts: Speech and Brain Mechanisms (Princeton: Princeton University
Press; 1959).
78
Evolution through Social Adaptation
2 per cent of the body’s bulk. It burns up a correspondingly great
amount of oxygen and sugar, which have to be fed to it con-
stantly.5
If the brain is an expensive superstructure for an adult to carry
around, it is even more of a burden for infants and children, who
have to be protected and fed longer than the young of other ani-
mals. At birth it has already reached 24 per cent of its adult
mass, whereas the whole infant body is only 5 per cent of its
adult body weight. At the age of three, the brain has attained 82
per cent of the adult weight and the body only 10 per cent.
When the child is ten years old, shortly before puberty, the brain
has attained 95 per cent of its adult volume, and from there on
it gains very slowly and very little, whereas the body grows
rapidly.6
In order to justify its carrying charges, any oversized and over-
fed organ has to have a selective advantage in the reproductive
life of the animal burdened with it, or its frequency will be kept
down by natural selection. This has been shown many times in
studies of other animals, the most conspicuous example, perhaps,
being that of antler size in the deer family. Putting it very simply,
there must have been a point in human history at which brains
came to be more effective than brawn in acquiring women. Other-
wise the brain sizes of various lines of fossil men would not have
increased during the Pleistocene. Just how the brainier men won
out is not known, except through analogy with living peoples.
Clever planning, self-control at the right moments, persuasive
talking, the exercise of leadership through language — these are
obvious possibilities.
The importance of brain size in relation to more complex social
behavior is suggested by comparisons with certain animals. Of all
the mammals, only the whales have larger and more complex
brains than man. The porpoise Tursiops truncatus, which is a
small and very bright species of whale, has a very complex brain
one third larger than ours, and a highly developed social life. In
5 C. F. Schmidt: The Cerebral Circulation in Health and Disease (Springfield,
111.: Charles C Thomas; 1950).
6J. H. Scott: “The Growth of the Human Face,” PRSM, Vol. 47, No. 2
(1954), PP- 91-100.
79
On the Antiquity of a Human Type of Society
it can be observed clear dominance relationships, and also al-
truism. Care, anxiety, and friendship between individuals have
been seen in the behavior of porpoises (as well as in that of
chimpanzees and some other primates).7 Furthermore, the por-
poises have possibly the most elaborate system of vocal com-
munication of all the nonhuman mammals.
On the Antiquity of a Human Type of Society:
the Beginning of Hunting
Before we can assume that the progressive increases in brain
size seen in the fossil record constituted, at least in part, an adap-
tation to the requirements of living together in a human society,
we must establish the antiquity of our basic social system, which
consists of a number of families living together and sharing food.
We can never do this absolutely — social structure is not a material
object that can be fossilized — but we can try to zero in on the
point at which it may have begun by following several lines of
evidence, including archaeological sequences, comparative ani-
mal behavior, and the social systems of living primitive peoples.
Let us begin with archaeology.
As previously stated, we may assume that the sharing of food
must certainly, because of the nature of the beasts eaten, have
begun with hunting, if indeed it had not already been practiced
earlier among food gatherers. We can gain some idea of when
hunting began by examining the camping sites at which fossil
men, or other manlike primates, lived, or at least made their tools
and ate.
The two oldest seem to be Bed I at Olduvai Gorge, Tanganyika,8
and Tell Ubeidiya in the Middle Jordan Valley just south of Lake
7 A. F. McBride: “Meet Mr. Porpoise,” NH, Vol. 45, No. 1 (1940), pp. 16-29.
McBride and D. O. Hebb: “Behavior of the Captive Bottle-nose Dolphin
Tursiops truncatus,” JCPP, Vol. 41, No. 2 ( 1948), pp. 111-23.
W. R. Thompson: “Social Behavior,” in A. Roe and G. G. Simpson: Behavior
and Evolution (New Haven: Yale University Press; 1958), pp. 291-310.
8 L. S. B Leakey: “A New Fossil Skull from Olduvai,” Nature, Vol. 184, No.
4685, pp. 491-3; and “Recent Discoveries at Olduvai Gorge,” Nature, Vol. 188,
No. 4755, pp. 1050-2.
80 Evolution through Social Adaptation
Tiberias in Israel.9 Both are Lower Pleistocene, and both were
discovered in 1959. The Olduvai camp contained a fossil manlike
primate which its finder, L. S. B. Leakey, named Z injanthropus,
and a second one, the so-called Olduvai child, both of which
will be described in Chapter 7. What is important here is that
crude stone implements as well as bones which showed signs of
being the remains of animals eaten on the spot were scattered
there. The tools were sharp enough to enable the hominid who
used them to cut skin, which he could not tear with his blunt
teeth nor soften with fire, which he lacked. But the bones
suggest that he had only begun to hunt: most of his quarry
consisted of small, slow-moving animals, like rats, lizards, snakes,
and tortoises, which can be caught by women and children. An-
thropologists call this category of animals slow game.
Leakey also found a few bones of the newborn and suckling
animals of large species of ungulates (hoofed mammals).
Whether this evidence places the Olduvai creature on the thresh-
old of life as a hunter is not certain; baboons have been seen,
by S. L. Washburn and others, to eat the newly born fawns of
impala. Eating newborn ungulates is hardly hunting, but it is a
gastronomic exercise that gives an animal a taste for fresh meat.
The Jordan Valley site contains tools, very fragmentary human
or humanlike remains yet to be described, and animal bones that
not only had been broken but also had been scratched with stone
tools. Most of the animals eaten seem to have been slow game, as
at Olduvai, but some appear to have been adult ungulates. All
the geologically later habitation sites we know of, in the higher
levels of Olduvai Gorge itself, in North Africa, China, and Europe,
indicate full-scale hunting.
Present evidence therefore suggests that true hunting, as op-
posed to the collection of slow game and baby animals, began, as
a way of life, sometime during the Lower Pleistocene, and we are
sure that in the Middle Pleistocene it was in progress. Speech
probably began with full-scale hunting, and a human kind of
social organization must have begun with speech.
9 M. Stekelis, L. Picard, N. Schulman, and G. Haas: “Villafranehian Deposits
Near Ubeidiya in the Central Jordan Valley (Preliminary Report),” BRCI, Vol.
9-G, No. 4 (i960), pp. 175-84.
The Mating Systems of Other Animals
81
The Mating Systems of Other Animals
Whether or not the members of an animal species enjoy
tools and speech, the social structure of the species is linked to its
mating system, which is further linked to factors of body size,
terrain, feeding habits, and climate, including seasonal change.
Small animals, like the familiar chipmunks, often live alone in
solitary burrows no more than 200 to 300 feet apart, individuals
of each species populating a neighborhood and dividing the
feeding grounds among them.
Each such animal has a home, in which it sleeps, and a home
range, in which it feeds or collects food.1 It will defend its home
against intruders of the same species, but its home range overlaps
those of other individuals, whose presence it tolerates in the com-
mon marginal areas. In these shadily defined territories infre-
quent and seasonal sexual contact takes place. As the offspring
are reared by the mothers alone, no elaborate social structure
arises and evolution through social adaptation is virtually non-
existent.
Larger animals, particularly hoofed and horned browsers and
grazers, tend to congregate in herds wherever grass and leaves
are abundant enough to feed many animals at once. Among such
animals which live on bulky foods, sexual activity consumes much
time, energy, and attention. In many species the sexes are sepa-
rated during most of the year, there being no reason, in the eco-
nomics of animal life, for them to be together. At breeding time,
which in many species comes but once a year, in a favorable sea-
son, the males and females come together and the males compete
for sexual rights. This period is necessarily brief because, being
preoccupied with sex, the animals are especially vulnerable to
carnivores. Most if not all females are covered, but some males
are left out. Those that impregnate the most does pass on their
genetic peculiarities, which consist largely of the traits essential
to success in courtship, such as larger antlers and stronger neck
1 F. Bourliere: The Natural History of Mammals (New York: Alfred A. Knopf;
1956), pp. 98, 220 f.
82 Evolution through Social Adaptation
muscles. This kind of selection enhances sexual dimorphism but
has little effect on other social capacities.
Yet another kind of mating behavior is found among mammals
that inhabit tropical forests, the special domain of primates.
Here no major seasonal change of climate or of food supply
makes any particular part of the year more favorable either for
birth or, by extension, for copulation. As man is a primate, other
primates will provide the most desirable comparative material for
study. In common with some of these primates, to be described
in the following chapter, man retains a very primitive sexual cycle
common to some of the simplest mammals, including marsupials
and insectivores. This sexual cycle is basic to our human social
structure. We have made the most, in this inherited trait as well
as in some others, of our lack of specialization.
The Sexual Behavior of Primates , Including Homo Sapiens
Members of most species of primates breed around the calen-
dar in roughly monthly cycles, which in the female include the
crises of ovulation and menstruation. Among the primates the
male is always ready for sexual activity whereas the female’s in-
terest is variable.
Two sets of physiological changes can affect her: differences in
libido during stages of the oestrual cycle, and the presence or ab-
sence of temporary genital swelling. Among some species the fe-
male feels an irrepressible urge for sexual attention at the time
of ovulation, pinpointing the proper moment for conception. Dur-
ing the rest of the monthly cycle she is either indifferent to sex,
or even resistant. Among other species the female feels equally
receptive at all times. In some of the species marked by a power-
ful urge on the part of the female during ovulation, she presents
an added stimulus. At this time her genitals puff and swell and
turn bright red, creating a conspicuous target that no male can
fail to recognize for the signal it is. Among other species that go
through the same cycle the aggressive behavior of the female at
this time is unaccompanied by such a display. In no case do we
find swelling without a marked increase of libido at ovulation.
Australoid subspecies: a Tiwi from Melville Island.
I
Mongoloid subspecies: a Formosan aborigine of the Bunun tribe.
II
Congoid subspecies: a Shilluk from the Sudan.
IV
Capoid subspecies: a Bushman woman from the Kalahari.
jilaal
V
Environmental adaptation: Allen’s Rule. Two Dinka girls from the Sudan.
VI
A Norwegian physiologist studying the cold tolerance of an
Alakaluf Indians foot: Dr. Kristian Lange-Andersen and
Lucho.
The social importance of fire: Bushmen of the Kalahari.
VII
Leadership in operation at a Tiwi funeral: the man on the
pole is directing dancers.
VIII
a. Common lemur
b. Ring-tailed lemur
c. Slender loris
d. Tarsius
Prosimians
a. Marmoset
b. Capuchin
c. Ornate spider monkey
New World monkeys
c. Brazza monkey
d. Patas monkey
Old World monkeys
XI
XII
b. Proboscis monkey
Old World monkeys
XIII
Orangutan
XIV
Chimpanzee
XV
Mountain Gorilla
XVI
The Sexual Behavior of Primates 83
The first kind of female behavior, the markedly variable, with
or without genital swelling, leads to a social structure in which one
male, whose libido is constant, serves a number of females in suc-
cession. Either he does this as a harem master, brooking no
rivals, or as a member of a mutually tolerant team of males that
take turns with female after female as their moments of high ex-
citement arise. The result is a choice of two social systems, a
harem or a club.
The second or relatively invariable kind of female behavior
creates the habitual association of one adult male, who is inter-
ested in sex every day, with one generally receptive female. The
result then is a third kind of social system, the monogamous
family.
According to Kinsey and his associates,2 59 per cent of Ameri-
can women interviewed reported that they enjoyed sexual inter-
course more at certain points of the oestrual cycle than at others.
Of these, only 11 per cent, or 6.5 per cent of the entire sample,
preferred it during the middle of the cycle, near ovulation. The
other 89 per cent, or 52.5 per cent of the whole, found it most re-
warding just before or after menstrual flow, or both. To what ex-
tent American women are typical of the human female of all
races and cultures is impossible to say. However, in primates in
general these characteristics are specific. Women of different
races are probably basically alike in this respect.
Although the human population of the world is growing at an
alarming rate, and we are having what is called a population ex-
plosion, nevertheless, of a good sample of the very women respon-
sible for this explosion, only 6.5 per cent seem to have felt more
passion during intercourse at a time useful for conception than
at other times. Also, the nonproductive added urge of half of the
women before and/or after menses helps them insure attention
from their husbands before and after a period of isolation, thus
re-enforcing a relationship between marital partners.
As far as reproduction is concerned, all the sexual activity that
2 A. C. Kinsey, W. B. Pomeroy, C. E. Martin, and P. H. Gebhard: Sexual Be-
havior in the American Female (Philadelphia: W. B. Saunders Company; 1953),
p. 608. Libido was measured objectively, by observing the amount of vaginal
secretion, as well as subjectively, by having questions answered.
84 Evolution through Social Adaptation
takes place among human beings, except at ovulation, is a waste
of time, energy, and attention. In addition, many human females
enjoy sex long after menopause. Nature is never wasteful, though,
and in fact the sexual behavior of the human female is oriented
more toward the maintenance of the social structure than toward
reproduction. It tends to create a family, an economic unit built
around the feeding, care, and education of children, and to secure
the continued interest of husbands. What is wasted in one sense
is gained in another. As we shall see in more detail later, this pat-
tern is also typical of the other primates whose females resemble
ours in this aspect of physiology.
The Beginnings of Human Society
Among animals that do not share food, and man is the
only higher primate that does, the function of the family, if any,
is to bring up the young ones to the time, or point of develop-
ment, at which they can fend for themselves. In different species
the age of parting from the mother varies. Among the apes the
youths are driven out of the family band at about the time of
puberty, not so much because they could not feed themselves
earlier, but because at that time they begin to arouse jealousy, in
the well-known Oedipus fashion, in their parents. The daughter
antagonizes her mother because of her father’s attentions and the
son antagonizes the father because of his advances toward his
own mother, or of the mother’s toward the son.3 So both son and
daughter are expelled, one at a time rather than simultaneously
since, as most primates have single births, the sons and daughters
will arrive at puberty at different times. This staggering of ex-
pulsions normally prevents mating between brothers and sisters
and encourages the mating of individuals of like ages simultane-
ously expelled from different familes.
At the time of expulsion the offspring, at first singly and then
paired, are strong enough and aggressive enough to block out
sleeping and feeding territories of their own and to defend them
against other family groups, including their parental households.
3 In a family of the harem type the “mother” can be any one of the wives.
Sexual Selection among Higher Primates 85
The motivation behind the parents’ behavior is primarily social,
but it has an economic side as well. Each newly weaned infant
will naturally stay with its parents as long as it can since finding
food where they do is easier for it than discovering new feeding
places in unfamiliar territory. But if all the offspring borne by
the couple were to remain with their parents indefinitely, there
would soon be a food crisis. The daily traveling range would have
to be widely extended, or some animals would go hungry. The
balance between territorial size and the number of mouths fed
can be maintained only by expulsion.
This composite picture of a prehuman primate society can serve
as a model for our ancestors before they became organized into
groups of mutually dependent families, either at or before the
onset of hunting. Whether or not it is a true picture depends
basically on how ancient the characteristic behavior of the human
female is — that is, how long she has been, as she is now, sexually
receptive at all times except during menses and not much more
aggressive at one time than at another.
A clue to this problem comes from the birth sex-ratio. Other
primates with our type of sexual behavior have a ratio of births
of about one male to one female. Those that live in harems or
clubs have more nearly three females to each male. But unfortu-
nately we do not know the birth sex-ratios of fossil men. We have
not found enough specimens of any population more ancient than
Sinanthropus, and even with Sinanthropus we cannot be sure of
the sex of each individual, usually fragmentary, specimen.
The chances are, however, that the ratio has been the same as
long as our record extends, for physiological matters of this kind
are conservative. This does not necessarily mean that all peoples
have always been monogamous, only that in most societies most
individual men are. In most societies exceptional men are polyga-
mous.
Sexual Selection Among Higher Primates
Among several species of higher primates a minority of males
seem to go through life without sexual experience, living as soli-
tary outcasts, or as neuters on the fringes of family groups. Among
86 Evolution through Social Adaptation
the majority some have more success in mating than others, and
the females, at least among chimpanzees, show marked prefer-
ences for individual males. Even though females also manifest
personality differences, however, all females normally receive
sexual attention. Among primates it is easier to be a female than
to acquire one.
Sexual selection, therefore, works particularly on males, and
whatever genes make a male a more successful lover than his
fellows can be expected to remain at a high level in the primate
pool. This may be why, among gorillas, males are twice as big as
females and it may help explain the extreme aggressiveness of the
males in many primate species.
Loving and intelligence do not necessarily go together, in apes
or men. But in the human evolutionary line, from the beginning
of hunting on, being bright has been an asset to a man in securing
the favors of women. He who brings in the most meat feeds the
most people, and they give him their daughters in order to insure
continued favors. He who handles weapons most skillfully against
wild beasts can turn them most lethally against his rivals in the
camp. An effective leader who can persuade others to work for
him can also outmaneuver his less well organized if more muscu-
lar rivals in the game of love.
Speech , Hunting, and Social Structure
Leadership and persuasion require thinking and talking,
and so do clever schemes. The threshold of becoming human
which our ancestors once crossed was largely the barrier between
communication by grunts, screams, facial grimaces, postures,
nudges, and bites on one side of the line, and articulate speech
on the other. Language not only made communication easier and
clearer, but it also increased its volume. We talk more than we
act, and, if we are wise, we think even more than we talk.
The study of speech is a whole world in itself, elaborate and de-
tailed, involving physiology, which we have discussed, as well as
psychology, history, and many other disciplines. It mirrors all
other subjects that concern man and his behavior. It is difficult
Speech, Hunting, and Social Structure 87
to imagine a world without speech, because imagination is im-
possible without it. We think in words. Words are what culture
is made of, and man alone creates and wields words and has
culture. Despite the importance of speech we know little about
its beginnings, for it was invented in the dim time about which
only specialists in human paleontology and flint archaeology
know, and it leaves no imperishable remains.
Whenever it started, once our ancestors had begun hunting,
speech was necessary. A hunter needs weapons. Weapons must
be made with tools, tools have to be made, and tool-making must
be taught. Beyond a certain level of technical skill, teaching re-
cpiires language. Hunting also requires planning, and planning
calls for speech elaborate enough to permit a group of men to
talk over, in the evening, what they intend to do the next day.
This is a much more advanced type of communication than the
common primate practice of uttering imperatives to signal im-
mediate action,4 which is all that the apes and monkeys can man-
age.
The social consequences of hunting need language too, even
more so than the planning and organization of the chase itself.
Except in big, usually annual communal hunts, in which women
and children beat the bush and drive game into the center of a
circle for the men to kill, hunting separates men from their
women. For two or three days and nights a married woman must
remain in or near the camp, exposed to the possibility of advances
from the old men and perhaps cripples left behind, and from the
boys too young to go hunting but old enough to be interested in
women. If the hunt is to be successful, if all are to eat meat, and
if the band is to retain its composition and integrity, these males
must leave her alone, and if any one of them should make the
mistake of approaching her, she must refuse him.
If this were not so, her husband would not go out at all, or if
he did, he would be so preoccupied when all his attention should
be concentrated on his task of finding, following, stalking, and
killing animals that he might fail. Even if his hunt were success-
ful, after his return to camp he would have discovered his wife’s
4 A. S. Diamond: The History and Origin of Language (New York: Philosophi-
cal Library; 1959). Also C. F. Hockett: op. cit.
88 Evolution through Social Adaptation
infidelity and there would have been a fight. The survival of any
group of hunters depends on the existence of rules governing the
behavior of women during their husbands’ absences, and on the
enforcement of these rules. Such rules cannot be formulated or
enforced without language.
In comparing human behavior with that of other primates we
must remember that a man can cover more ground in a day than
a monkey or ape, that he can carry food and water, that with a
stick he can dig roots that lie too deep for the fingers of monkeys,
and that he can feed off a larger territory. All else being equal,
more human individuals can live together in societies than can
other large primates, including those that have our kind of fe-
male sexual physiology.
When, therefore, people began sharing food, it was no longer
economically necessary to expel both the boys and the girls
from the group at puberty. A much more effective system was
for either the boys or the girls to leave the parental domain,
marry into other households, and live with their in-laws. Such an
arrangement tended to foster peaceful relations between neigh-
boring bands, under cover of which gene-flow could extend over
a wide area within natural boundaries.
Within individual households, the older men need no longer
be killed, driven off, or reduced to a servile state once their
strength had begun to ebb. With the power of speech and a long
period of dominance behind them, they could persuade the young
men to feed them, and might even bluff them into allowing them
to have the most desirable women. When the band grew too big
for the territory it inhabited, it could simply split under individ-
ual leaders, and the pioneer half could set out to find and exploit
a new territory of its own. If, on the other hand, a band grew too
small for efficient operation, it could combine with a neighboring
and related group.
Ritual, Language, and the Rites of Passage
The association of several families in a band brought a
dozen or more children together. Now they could play games in
Ritual, Language, and the Rites of Passage 89
groups large enough to permit them to sort themselves out into
leaders and followers and to learn co-operation. Also, the older
children then had a chance to teach the younger ones. By the time
a child reached puberty, he or she would have learned more than
could have been possible had the families lived apart, and in par-
ticular the children would have begun to learn how to get along in
groups.
But once the children came to puberty, changes in endocrine
balance exposed them to new and violent stresses, which required
a special schedule of indoctrination, if order was to be main-
tained in the camp. The young men had to be segregated in
classes, usually recruited from several neighboring bands. They
had to be sent out into the wilderness to fend for themselves, with
restrictions on certain foods to make the apprenticeship harder
than real life; taught obedience by the shock method through the
appearance of old men disguised as supernatural creatures; and
carefully instructed in the proper behavior toward women.
When this “probationary” period was over, they were readmitted
into the company of their parents and other relatives as partial or
full-scale adults.
Without a puberty ceremony it is difficult to see how the transi-
tion from expulsion to incorporation of the young could have been
managed. And without language such a ceremony would be im-
possible. We can be confident, therefore, that language goes back
at least as far as this major change in human social organization.
With the awareness of natural processes that language brought,
along with a keen observation of every phase of plant and animal
life on which human life itself depended, came a full realization
of the inevitability of death. People began to generalize and to
reason, using the materials at their command as symbols, and
building up imaginary worlds of spirits that controlled plants and
animals, and of spirits of dead people. These elaborate structures
were necessary by-products of man’s growing intelligence. He
needed them to allay the fears that his new knowledge of the
world of the senses brought him, including the inevitability of
death. With spirits to help him and an afterlife to look forward to,
he could tolerate fear, and create ceremonies for other crises
beside puberty, including the crises of birth, of death, of changes
90
Evolution through Social Adaptation
in the food supply brought by the cycle of seasons, and of changes
in the routine of interpersonal relations consequent on shifts in the
seasonal round of activities.
Homo was becoming a more and more sensitive animal, in-
creasingly vulnerable to social disturbance as he came more and
more to perceive and use the forces of nature. Individuals who
could learn to speak easily had an advantage over those who
could not speak at all. Once the use of language had begun, selec-
tion in favor of facile talkers must have been an important factor
in the stages of human evolution that followed.
The Discovery of Fire and the Conversion
of Energy into Social Structure
Three innovations that had to be learned — walking erect, tool-
making, and speaking — prepared our ancestors to organize them-
selves into bands of families that hunted, shared food, and con-
ducted ceremonies together, but it is highly questionable that
these three were enough to make a human social structure pos-
sible. One more ingredient was needed. That was the use of fire.
As I have pointed out elsewhere,5 human beings convert energy
drawn from outside their own bodies into social structure, and the
greater the amount of energy consumed, all else being equal, the
more complex the social structure. Such a use of energy increases
the physical efficiency of people, individually and as groups, to
such an extent that the time spent in obtaining, processing, and
eating food is reduced. At the same time, economic activities are
shared and divided among the members of the group. The
division of labor based on sex, age, and kind of activity which is
thus made possible fosters further efficiency. The new relation-
ships between individuals and groups so created acquire more
and more social complexity.
Fire has four basic uses: frightening off predators, keeping peo-
ple warm and dry, cooking food, and providing a spatial nucleus
or center for the home territory of a group of people. Here they
can sit at night, warm and secure, seeing one another’s faces in the
5 C. S. Coon: The Story of Man, p. 64.
9i
The Evidence of Living Food-Gathering Societies
firelight, talking over what they did during the day while they
were separated, acting out scenes of the hunt, planning for the
next day’s adventures, discussing matrimonial prospects, and gen-
erally getting to know one another so well that friction can be kept
at a minimum. They may also dance by firelight, and conduct
ceremonies. It is difficult to see how, without fire, human society
could have risen much above the level of that of baboons.
If we ignore the Australopithecines, who were probably not
full-scale hunters but collectors of slow game, we may state that
fire is as old as the oldest undisturbed sites of the genus Homo who
lived in chilly climates. The oldest men in the Far East, the Sinan-
thropus population of Choukoutien, had it 360,000 years ago.
Evidence of fire has been found at Swanscombe, England, in the
same level as the Swanscombe skull, which is over 250,000 years
old. The only older specimen in Europe, the Heidelberg or Mauer
jaw, was taken from a secondary deposit in a gravel pit. None of
the early remains from Java were found in habitation sites. The
oldest found to date in Africa south of the Sahara is only 40,000
years old. Despite the careful excavation of several older undis-
turbed habitation sites in East Africa, no earlier trace of fire has
been found there.
The Evidence of Living Food-Gathering Societies —
the Australian Aborigines
Tucked away in odd corners of the earth are several hun-
dred tribes and other population units of people who still live by
hunting and gathering. Technologically they represent every level
of competence discovered by archaeologists. A few make and use
stone chopping tools, others manufacture simple flakes, and so on
up through the chronological list of archaeological implements to
the threshold of metal. Their housing ranges from simple leaf
windbreaks to elaborate wooden buildings, and their clothing
from complete nudity to the world’s most efficient arctic suits.
From our present point of view, food gathering is not a single
way of life. The Indians of the northwest coast who harvested
salmon and hunted whales attained a high cultural status without
92
Evolution through Social Adaptation
agriculture; and the circumpolar peoples, from Lapland to Green-
land, were able to live in an otherwise uninhabitable world only
by the exercise of great ingenuity. Neither of these groups repre-
sents the stage of cultural evolution we are seeking. To find it we
must turn to the marginal refuges of the Southern Hemisphere, to
the forest recesses of the Oriental and Ethiopian regions, and
particularly to Australia.
Australia contains the world’s most archaic mammalian fauna,
and it also harbors the world’s largest assemblage of archaic hu-
man beings. However, neither man nor his prey has been there
very long. The monotremes, unique to that continent and New
Guinea, are unknown before the Pleistocene. The marsupials ap-
peared in the Pliocene, and man toward the end of the Pleisto-
cene, about 11,000 years ago, or a little earlier.6
The monotremes apparently evolved locally from reptilian an-
cestors. The marsupials entered the Australian faunal region over
an unknown path from the New World, and Australoid people
arrived from Indonesia by island hopping while the seas were still
low, crossing Wallacea from the Sunda to the Sahul shelf, prob-
ably on flimsy rafts and canoes of types still made in modern
times. Linguistic theory (see Chapter 1, p. 5) supports both a
late date of under 20,000 years ago and an invasion or series of
invasions from a single source, because all Australian languages
belong to a single family.
Many studies have been made of Australian social systems but
most are too specialized for our purpose, nor are they organized
from a biological viewpoint. They overconcentrate on theoretical
marriage regulations and give too few case histories and statistics.
In general, they tell us that Australian aborigines live in house-
holds of a few families, each in its own hunting territory, and that
from time to time a number of related households meet to conduct
ceremonies jointly. These may include the initiation of a new class
6 N. B. Tindale: “Ecology of Primitive Aboriginal Man in Australia,” in A.
Keast, R. L. Crocker, and C. S. Christian: “Biogeography and Ecology in Austra-
lia,” MB, Vol. 8 ( 1959), pp. 36-51. Tindale gives a Carbon-14 date of 8,700 ± 120
years ago (about 6,750 b.c.) for a site at Cape Martin, southern Australia, contain-
ing the Tartangan culture, which was preceded by the Kartan culture. The Kartan
antedated the rise of the sea level at the end of the Pleistocene, about 10,000
years ago. The laboratory number of the date given above is NZ-69. For an ex-
planation of this symbol, see note on page 311.
The Evidence of Living Food-Gathering Societies 93
of boys and marriage negotiations. In these meetings the older
men play a dominant role, just as they do in most other human
societies.
The collection of households that meets on such occasions is, in
effect, a breeding isolate in the zoological sense. Within its con-
fines rules of various degrees of complexity specify which men are
eligible to marry which women, because of their membership in
certain segments of the total population. They acquire this mem-
bership by descent. Usually a man can marry only women from
a group more distantly related to him than others. Thus only a
fraction of the women are theoretically available to him, never
more than half, and sometimes as few as one thirty-second. Within
these limits he can have one or more wives. If no spouse is avail-
able at all, the rules can sometimes be stretched to include some
other women almost equally distant in kinship. These rules serve
to split up the breeding population into a number of smaller
isolates that rarely intermarry.
As a man can have several wives at any one time and a woman
can be impregnated by only one husband at a time, a man can
have more children than a woman can. Natural selection thus
tends to favor characteristics borne by the male. At first glance it
would seem that disparity in reproduction among males would
have no evolutionary value, being based on accidents of the birth
ratio, but this is not the case. A dominant male can manage to
have one or more wives by manipulating the marriage system,
and a less aggressive, less clever, or less competent male may be
left out. As the traits of personality that give some men more
women than others are inherited, selection in favor of these traits
must occur. On the other hand, women are the prizes of masculine
competition and, although some scheming among women also
takes place, no woman is sexually neglected who is still able to
bear children.
In surveying the literature on Australian social systems, we are
soon struck by the great differences in age between husbands and
wives. An old man may be married to two teen-age girls, and a
younger man to a withered crone. As they can rarely count to ten
and have no measure of the passage of years they do not know
how old they are, and are in effect as young as they look and feel.
94 Evolution through Social Adaptation
If a man has to wait twenty years for a wife, it is not twenty years
to him, but simply a long time.
The Archaic Society of the Tiwi
An Australian tribe that has been intensively studied
recently is the Tiwi, who inhabit Melville and Bathurst Islands.7
They number about 1,000 persons concentrated near three white
settlements, with 600 at a Catholic mission on Bathurst, 50 out-
side ( but not part of ) a government half-caste station at Garden
Point on Melville, and 150 at a government station at Snake Bay
on Melville, in addition to 150 who are working at Darwin.
Within the lifetimes of the older Tiwi, their islands were di-
vided into ten “countries,” each occupied by a number of house-
holds consisting of one or more families, each in its own hunting
and food-collecting territory. As families grew, split, shrank, or
combined, so did both the territories and the countries. But the
surface of the islands, comprising about 3,000 square miles, was
open to all of them when they met to take part in ceremonies, par-
ticularly funerals. They spoke one language and were in a very
loose sense a people.8
The Tiwi went naked and built flimsy shelters to serve as sun-
shades and as protection from heavy rain. Their only cutting tools
were an all-purpose clam shell and a flaked stone ax, poorly
hafted and reminiscent of a chopping tool.9 A few crude stone
flakes were used solely for gashing foreheads at funerals.
Melville and Bathurst Islands are well forested and rich in both
7 C. P. Mountfort: The Tiwi, Their Art, Myth, and Ceremony (London: Phoenix
House; 1958).
J. C. Goodale: “Alonga Bush, a Tiwi Hunt,” BUM, Vol. 21, No. 3 (1957),
PP- 3-36.
Goodale: “The Tiwi Dance for the Dead,” Expedition, Vol. 2, No. 1 (1959),
PP- 3-13-
Goodale: The Tiwi Women of Melville Island, North Australia (Philadelphia:
University of Pennsylvania Ph.D. dissertation; 1959).
C. W. M. Hart and A. R. Pilling: The Tiwi of North Australia (New York:
Henry Holt & Co.; i960).
8 Although the Tiwi still practice many of the customs summarized here, they
have abandoned others. I am using the past tense only for continuity.
9 However, better axes with pecked and ground surfaces were found on the
beach at Snake Bay. Their age is unknown.
95
The Archaic Society of the Tiwi
animal and vegetable foods. Wild yams, wallabies, and opossums
can still be had on land and shellfish are always available at low
tide. The surrounding sea and its estuaries contain turtles, croco-
diles, and fish, and fresh-water swamps at the heads of creeks pro-
vide food for wild geese. Every year the Tiwi burned over the
landscape to keep down the undergrowth which impedes hunting.
All vegetable foods belonged to the women, who dug and col-
lected them. Animal life above the ground, particularly wallabies
and opossums, belonged to both men and women and both
hunted it. Both could also collect shellfish. The beasts of the sea,
including fish, and the fowl of the air belonged to the men alone.
A young man who was a poor or indifferent hunter, and lacked
a pleasing personality, could kill enough marsupials to feed him-
self, but he could not bring in the quantities of meat obtainable by
killing sea turtles, crocodiles, and geese. All these animals were
hunted by teams, and a boy had to be invited to join such a team.
To catch sea turtles and crocodiles, the men traveled by canoe.
Usually the owner of the craft paddled in the stern, a boy bailed
amidships, and another man stood in the bow with his spear. Only
one man made the kill, but all three shared in the meat. Sometimes
geese were killed by solitary hunters, but usually men teamed up
to cover a greater area. The hunters would spread out along the
bank, evenly spaced, to await the geese, which flew over in small
flocks. Usually only one man was in range, and no one knew which
man this would be.
There was an element of danger in going out in canoes, for the
men could drown or be eaten by crocodiles. In goose hunting the
accent was on good marksmanship and reliability. The brave,
skillful, obedient young man accepted by his elders as a hunting
partner was able to feed several persons with his share of the meat.
Thus he had a quid pro quo for obtaining wives, one which held
both economic value and prestige. The indifferent hunter who was
not wanted as a teammate was no more useful as a food provider
than an indiff erent woman.
Good hunting and good partnership, however, were not the
only roads to popularity and prestige. The Tiwi put great store in
aesthetic achievement. A poet who could compose and sing a new
and popular song, a dancer who could create a novel routine, and
96 Evolution through Social Adaptation
an artist who could paint stimulating designs on their ultramod-
ernistic-looking funeral poles also rose to the top of the Tiwi social
ladder and to the intimate companionship of those who dispensed
matrimonial largesse.
Once the superior young man — who had to be bright to be
superior — had passed the age of thirty and had secured several
wives of various ages, he could leave the provision of staple foods,
yams, and marsupial flesh to them and could concentrate on sup-
plying the prestige foods from the sea and air to a wider circle as
he connived at a game of competitive prestige to become more
uxorious than ever. When he had finally become too old to hunt,
he would have had plenty of people to feed him. In Tiwi society,
therefore, a combination of hunting skills, good teamwork, cour-
age, and artistic and political competence gave superior men the
greatest procreative opportunities, and some men were sloughed
off from the gene pool through incompetence on any or all of these
counts.
Although greatly condensed, this sketch of Tiwi society is ac-
curate enough to demonstrate the reality of social selection. A
little detail may further clarify the point. When a boy arrives at
puberty, he is usually promised the future daughter of a girl his
own age who herself has been spoken for since before her birth.
This girl, the boy’s future mother-in-law, now becomes the wife
of a man at least thirty years old who has been waiting for her all
her short life, and who has been a food provider and a constant
visitor to her parental household. The future husband of her as yet
unconceived daughter, whom she may or may not bear, now also
becomes a food contributor and visitor to this growing house-
hold. This boy now busies himself providing his future mother-in-
law with meat while her husband is still feeding her mother. The
better the boy is at obtaining meat the more he pleases other men
whose daughters he might also be able to support on the same
promissory basis. Such older men will be likely to invite him to go
hunting with them.
However, even the most successful man cannot contract for just
any unborn girl. Tiwi society is divided into four phratries,1 sub-
1 A generation ago one of these split into two, making a total of five, but this
is unimportant for present purposes.
The Archaic Society of the T iwi 97
divided into a total of twenty-one clans, with two to six clans to a
phratry. Ordinarily, men of phratry A take their wives from any
clan of phratry B, and vice versa; C and D similarly exchange
wives." This produces in effect two breeding isolates in Tiwi so-
ciety and limits the choice of wives to one out of every four girls.
In Tiwi as in other societies, men tend to die younger than
women, and because of the age differential at marriage, Tiwi hus-
bands often die long before their wives, leaving one or more
widows. These women, including also young girls still living with
their parents and others yet to be born, have to be remarried or
reassigned. Often the older widows go to the men of thirty or so
who might otherwise have to wait ten or twenty years or even
longer for their assigned brides to be born and to reach puberty.
At this point, through their connections with their elderly
wives’ offspring, they can sometimes wangle promises of other
young brides or pick up young widows. Cleverness in manipulat-
ing marriages was as important as skill in hunting sea animals and
waterfowl; indeed they were often related.
According to Hart and Pilling, some Tiwi men had over twenty
wives. They cite one man who at about age thirty started with two
elderly widows and at sixty-six had had six widows and fifteen
young wives, of which total three widows and five young wives
had died. His youngest wife, not yet nubile, was still with her
father. This left him twelve wives in residence at one time.
Those were the good old days. In the pallid present, ridden
with white men, things have changed. In the Snake Bay colony
sixty men have sixty-six wives, and no man has more than three.
Out of fifteen young men between twenty and twenty-nine, seven
are already married. In the old days they would still be waiting.
Five men between fifty and seventy are still bachelors and prob-
ably always will be. Four are the conventional marital failures
specified above. The fifth is an excellent hunter of solitary tem-
perament who never wanted to be married, and wasn’t.
According to local opinion three of a total of nine men of the
senior age group, in their sixties and over, are “big operators,” at
the top of the pile. In the next age grade, from about forty to sixty,
2 The rules are far more complex than this. I have simply summarized what
happens biologically.
g8
Evolution through Social Adaptation
five of twenty-five are the social elite. These eight dominant men
out of the thirty-four adult males of their combined age group, or
13 per cent, have sixteen of the thirty-six wives in the group, or 44
per cent; and twenty-eight of its forty-eight children, or 58 per
cent. No doubt others coming along nicely in the age group under
forty will do equally well later, if Tiwi culture persists.
Tiwi society does not conform to any other primate model, and
there is no reason why it should. In some ways it resembles the
monogamous, small territory system; in others, the harem system.
Like other primates, the Tiwi quarrel over women, particularly
when the young, libidinous men become impatient waiting for
wives. Young women, bored with their ancient husbands, seek
amours outside, and these affairs sometimes lead to blows; eyes
are knocked out and flesh wounds are inflicted with throwing
sticks and spears. Ordinarily a young man being punished for
sexual encroachment will let himself be wounded rather than kill
the venerable elder whom he has cuckolded. One man in his
fifties, the husband of three young wives, is kept busy fighting off
their paramours. As he has punished some of them more than their
offenses seemed to warrant, he has lost face and several times has
been exiled from Snake Bay by his peers. Their civilized attitude
is a far cry from the simpler reaction of harem masters among our
lower primate kin. The struggle over sex is still present in Tiwi
society, but it is in a new balance. Men who can create fine poetry,
dances that rival the ballet, and art that fills the moderns with
admiration do as well in the marriage game as bullies, if not
better.
Tiwi society is undeniably archaic. The Tiwi lie on the fringe
of a marginal continent; they are the most marginal of marginals.
They have never had spear throwers, stone-tipped spears, boomer-
angs, circumcision, or other elements of “advanced” Australian
aboriginal culture. Physically they are also archaic full-sized hu-
man beings with a plethora of heavy brow ridges and big teeth,
and brains of only moderate size. They have had the fortune to be
preserved in a geographical paradise in which an early and agree-
able form of human life can be led by healthy people without too
much effort, and they have the sophistication of participants in a
culture that has long since “arrived.” Having no sense of inferi-
On Comparing the Cultures of Living Food Gatherers 99
ority to the white man, they look on us busy overdressed people
with an air of amused and kindly tolerance.
Evidence suggests that natural selection is still taking place
among the Tiwi. Its function is to keep them human, to maintain
the ratio of genes that contribute to a civilized life in a stable en-
vironment and to keep down the ratio of genes that lead to anti-
social or solitary behavior. The population meanwhile remains
stable as a result of the mating of old men with young girls, which
is relatively unproductive; abortion practiced on the young
women to space their children; and probably through other bio-
logical forces of which we are not yet fully aware.
Had the Tiwi become extinct several centuries ago, there
would be nothing much — in the physical and cultural remains
archaeologists might unearth — to indicate the heights of art this
people had reached, the fun they had had, or the sophistication
of their way of life. Caveat excavator. When we come to talk
about the Neanderthals and other early folk, let this be borne in
mind.
On Comparing the Cultures of Living Food Gatherers
and Those of Fossil Men
The Tiwi are, of course, only one of several hundred surviving
food-gathering peoples, but they are particularly useful for pres-
ent purposes because, with most of the other Australian aborigi-
nes, they represent the survival, with little change that we can
detect, of a cultural level found elsewhere 70,000 to 100,000 years
ago. Most of the other food gatherers that we know about, out-
side of Australia, either are dwarfs, like the Negritos, Andama-
nese, and African Pygmies; or they are pedomorphic, like the
Bushmen; or they live in an advanced Mesolithic type of culture,
like the California Indians; or, like the Veddas of Ceylon, they
have long been trading with food-producing neighbors. Some are
both dwarfs and traders.
Each of these qualifications weakens comparisons that we may
try to make between such living food gathers and fossil men. We
have no fossil dwarfs or pedomorphs. The Mesolithic began a
mere ten or twelve thousand years ago, long after the time we are
MR. MICH EL PLOTKL *
1240 LENOX AVENUE
MIAMI BEACH 39, FLORIDA
100 Evolution through Social Adaptation
interested in. Trading relationships between peoples of different
cultural levels constitute a cultural form of the biological process
known as symbiosis, and symbiosis is usually accompanied by loss.
In a parasitic animal whole organs may be lost. In a parasitic cul-
ture whole procedures, such as tool-making, and certain rituals
may be lost, and the marriage system may be affected. Also, when
food gathers trade with villagers, exchanging forest products for
tools and luxuries, some of the more adventurous among the
young food gatherers may leave home to join the culturally more
advanced population, and this drainage, through selection, can
genetically impoverish those left behind.3
Population Size Among Food Gatherers
The T i w i population of about 1,000 persons is relatively large
for Australia. Birdsell 4 finds an average of about 530 persons per
breeding unit at the time of their first encounter with whites. This
unit, called tribe by Australian anthropologists, is the group of
related bands and households that come together at least once a
year in time of plenty for ceremonies, initiations, matchmaking,
and merrymaking. Krzywicki 0 has divided 123 Australian tribes
into size groups, as follows. Seventy, or 57 per cent, had under 500
persons; 37, 01 30 per cent, had between 500 and 1,000; 12, or 10
per cent, between 1,000 and 2,500; and only four, which were
probably confederacies and not breeding units at all, had over
2,500.
In the Andaman Islands the breeding unit was about 350 per-
sons; 6 among the root-gathering Kadars of the Cardamon Hills in
India it is 566; 7 and among the /Kung Bushmen 8 of the Kalahari
J. Emperaire and A. Laming: The Last Fuegians,” Diogenes, No. 8 (1954)
PP- 37-68.
4J- B. Birdsell: “Some Environmental and Cultural Factors Influencing the
Structuring of Australian Aboriginal Populations,” AN, Vol. 87, No. 834 (1953),
pp. 171-207.
5 L- Krzywicki: Primitive Society and Its Vital Statistics (London: Macmillan &
Co.; 1934), PP- 171-207.
6 A. R. Brown: The Andaman Islanders (Cambridge: The University Press;
1922).
7 U. R. Ehrenfels: Kadar of Cochin (Madras: University of Madras; 1952).
8 The symbol / as used here indicates a Bushman click.
Population Size Among Food Gatherers 101
around 750.9 In north America Krzywicki tabulated 232 food-
gathering tribal populations. One hundred tribes, or 43 per cent,
were under 500 persons; 63, or 27 per cent, were between 500 and
1,000; and 69, or 30 per cent, were over 1,000. These last were all
from the northwest and included many technologically advanced
people who harvested annually migrating fish and mammals.
Many were confederations, each including several biological pop-
ulations.
This statistical exercise shows that the ecology of food gather-
ing is the same nearly everywhere. The requirements of hunting
and collecting keep the number of people who live near enough to
one another to breed as a unit within about 500 or 600 individuals.
We may suggest, but we cannot prove, that most of the fossil men
whom we shall presently study lived in populations of this size or
even smaller. There is no logical reason why their populations
should have been larger, at least in the earlier periods.
In any group of 550 persons, if the children, the aged, and the
infertile adults are excluded, there would be less than 200 breed-
ers, of whom less than 100 would be males. As most of the living
food gatherers observe some kind of marriage regulation compara-
ble to that of the Tiwi, which splits them up into subgroups, the
actual breeding units of some of the early populations may have
been equally small. If our analogies are correct, human beings
must have lived in small populations for a very long time. A string
of small populations covering a continental region, with natural
selection taking place in different populations, and some gene
flow over the borders, is just what would have impelled the evolu-
tion of races during the Pleistocene.
Let us not forget, however, that peoples who harvest fish and
mammals in large numbers can have breeding units of over 1,000
persons. During the Late Pleistocene in Europe the Upper Paleo-
lithic peoples were killing reindeer in such numbers that they
could select the animals most desirable in age and sex and let the
others go. One mammoth feeds many mouths, and the mammoth
hunters of central Europe killed so many of these giant animals
that they were able to stack the bones in piles, each kind of bone
9Loma Marshall: “Marriage Among the /Kung Bushmen,” Africa, Vol. 29,
No. 4 (1959), PP- 335-65-
102 Evolution through Social Adaptation
to its own heap. It is not unlikely that these hunters lived in com-
munities of over 1,000 and that the same was true, later on, of the
Mesolithic and Early Neolithic fish trappers who inhabited the
lower banks of such rivers as the Elbe and Huang Ho.
Systems of Mating Among Food Gatherers
The T i w i are organized in monogamous or polygynous house-
holds— mostly the latter. The same is true of most Australian
tribes. Murdock has listed the matrimonial systems of 564 dif-
ferent peoples of the world, 88 of which qualify as food gatherers.1
I have added one, the Kadar,2 and changed one, the /Kung
Bushmen,3 on the basis of later information. The total is now 89.
Seventy-three of them, or 82 per cent, are polygynous; 15, or 17
per cent, monogamous; and one, the Kadar, is both polygynous
and polyandrous. Among Murdock’s food producers, 71 per cent
were classed as polygynous, 25 per cent as monogamous, and 4
per cent as polyandrous. It is clear, then, that man prefers po-
lygyny when this form of mating is possible, although most men
have only one wife. This is true of peoples of all levels of culture
from the Tiwi to the Turks, living in every continent and climate,
and belonging to all human subspecies. We can assume that it
goes back a long way in our life as a genus. Thus, the ability to
obtain more than one’s share of women may have been a factor in
human evolution for a long, long time.
The Longevity of Fossil Men
A remarkable feature of Tiwi society is the presence of a
number of old men past their muscular prime who occupy posi-
tions of prominence. Out of sixty men at Snake Bay, twenty-two
1 G. P. Murdock: “World Ethnographic Sample,” AA, Vol. 59 (1957), pp.
665-87.
2Ehrenfels: op. cit.
3 Lorna Marshall: The Kin Terminology System of the /Kung Bushmen,”
Africa, Vol. 27 (1957), pp. 1-25. Murdock listed the Bushmen as monogamous.
Marshall, however, finds some to be polygynous.
The Role of Isolating Mechanisms in Human Evolution 103
were over fifty and five were believed to be over seventy. These
figures do not indicate the proportion of such graybeards to the
total number of people born in a generation; some of the younger
men, we must remember, are at Bathurst and Darwin. Probably
the ratio of males who lived to that advanced age is low. Still, the
figure is impressive, particularly in view of the life expectancies of
many peoples today. Our question is, did any men live to be old
in the remote past?
The answer is yes, but not many. Of the twenty-five individuals
of all ages represented by the Sinanthropus bones unearthed at
Choukoutien, two were over fifty years old.4 Of a compilation of
thirty-four Neanderthals from Europe, two were “advanced in
age but not senile.” 5 In my opinion, the Sinanthropus remains
show that as early as 360,000 years ago some peoples had attained
a level of social organization in which men of fifty, who had
passed their physical prime, were tolerated, if not fed, by their
juniors. Later on, 70,000 to 45,000 years ago, the Neanderthals
definitely fed old and crippled men. La Chapelle aux Saints, the
most famous French Neanderthal, although not much past his
forties, was toothless and crippled by arthritis. Shanidar 1 was
born with a withered right arm, part of which was later amputa-
ted, yet he was well over forty when he was killed by a rockfall in
a cave. These aging cripples were being fed; and anyone who
feeds middle-aged cripples lives in a human type of social struc-
ture.
The Role of Isolating Mechanisms in Human Evolution
Two biological problems are central to the theme of this
book. ( 1 ) How did the subspecies of man become differentiated?
(2) Why did they not become separate species? In other animals
related species occupying the same territory (i.e., sympatric spe-
cies) or adjoining territories are kept apart genetically because
their members do not breed together, whether or not fertile off-
spring could be produced if they did. The biological mechanisms
4F. Weidenreich: “The Sinanthropus Population of Chou Kou Tien,” BGSC,
Vol. 14, No. 4 ( i935)> PP- 427-61 (also CMJ, Vol. 55 ( 1939), pp. 33-44).
5 H. V. Vallois: “La duree de la vie chez l’homme fossile,” CRAS, No. 204
(i937), PP- 60-3.
104 Evolution through Social Adaptation
or procedures that prevent interbreeding are called isolating
mechanisms. These take many forms.
Among certain invertebrates sexual relations between species A
and B are prevented by the fact that the genital organs of a male
of species A will not fit into those of a female of species B. Among
amphibia, such as tree toads, the pitch of the mating call may be
critical. Each species has its own special locus, or loci, on the sonic
scale, and males and females of a given species reach each other
by following these calls. Mammals also have specific calls, and in
the case of the moose, the sound of a female’s urine dropping in
the water of a swampy lake will rouse the bull’s libido to fever
pitch, whereas it would leave a male deer unimpressed. On the
whole, however, most isolating mechanisms in land mammals in-
volve the sense of smell, which is also vital to them in marking
out their territories.
Isolating mechanisms can arise only in isolation. Once two re-
lated subspecies have acquired different ones, they can meet with-
out interbreeding, and have speciated.
Such mechanisms do not exist in all kinds of animals. Fish that
spread their milt and eggs broadcast in the sea obviously do not
have them. Others that breed in special places at special times, do.
Large animals that have only one species to a genus do not ordi-
narily need them. In the case of man, we have modern evidence
that Mongoloids, whites, Negroes, and Pygmies each finds the
odor of the next one in the series unpleasant, but this olfactory
barrier (based on the number of apocrine glands) has not pre-
vented mixture between any two of these groups. What retards
mixture in modern societies, as in India, South Africa, and the
United States, is something else: there peoples of different races
have been brought together by historic or late prehistoric inva-
sions and kept apart by an ethnic division of labor probably un-
known to simple food gatherers. The barriers which separate these
racial and ethnic isolates are probably all products of the last
8,000 years of human life, that is, they are the fruit of technology,
which has permitted races brought together artifically to remain
apart longer than they could have done on a food-gathering level,
and longer than they may be able to do once our space-age world
culture becomes thoroughly homogenized.
The Role of Isolating Mechanisms in Human Evolution 105
Whether clearly differentiated subspecies or closely related and
potentially interfertile species which have been artifically juxta-
posed will remain genetically isolated depends to a certain extent
on whether or not both sexes are present in each population. For
example, at about 1907 a herd of wild mouflon that had been
brought from Corsica and Sardinia was released on Lambay Is-
land, Dublin County, Ireland. A domestic herd was also grazing
there, and the island was unfenced. These two kinds of sheep are
interfertile, and are either closely related species or well-differ-
entiated subspecies — it is hard to tell when one kind is domestic.
In zoos the two do not mix, nor did they on Lambay Island as
long as there were both males and females in each herd. But
owing to shooting and other causes, the mouflon herd declined to
two rams and one ewe. The rams remained faithful to their consort
until her death. Then the rams joined the domestic herd and in
one season sired twenty crossbred lambs that were fertile and
were absorbed into the domestic herd.6
Modern men behave much like the mouflons, but in reverse.
Sailors and explorers, whose wives have been left at home, mate
freely with native women of all races; but when the settlers follow
with their wives and children, race mixture is usually forbidden.
Some of the very soldiers, sailors, and marines who nearly created
a new race in the Pacific Islands in World War II are opposed to
the mingling of races in their native states. This is not incon-
sistency— it is simply biology.
But the races of man evolved long before modern technology
made exploration or colonization possible. Far more pertinent to
the subject of this book are the age-old systems of mating prac-
ticed by living food gatherers. In this respect food gatherers differ
from other animals in that food gatherers consider their marital
rights as property. A man who has many wives is a man of pres-
tige. A stranger who visits the camp and is considered important
enough to be sent home happy is loaned a woman. Many white
men have been so accommodated. A temporary exchange of wives
may be part of the peacemaking ceremony between tribes that
have been fighting and are reconciled. If one tribe defeats the
6J. A. F. Roberts: “A Geneticist’s View of Human Variability,” in P. Mason,
ed.: Man, Race, and Darwin (London: Oxford University Press; ig6o),pp. 48—55.
1q6 Evolution through Social Adaptation
other, the vanquished men may be slaughtered, but their women
and young children will be taken as prizes. Many examples of such
behavior can be cited. They are the common grist of anthro-
pology.
But how far back does this kind of sexual behavior go? It is
universal in Homo sapiens, including the Australians. Is it fair,
then, to assume that it is at least as old as our species and nearly as
old as speech? Because this mating system ensures gene flow
wherever populations meet, and because isolated populations —
the Australians, for example — have not been alone long enough
to speciate, we can tentatively consider this system as one reason
for our failure to develop watertight isolating mechanisms, and
for our unity as a species.
Adaptation to Crowding:
A New Theory of Evolution hy Succession
Animal species vary greatly in the number of individuals
that can live together and tolerate one another’s presence. Some
kinds of insects live without friction in hives and labyrinthine hills
in the hundreds and thousands. Some fish swim in schools, some
birds fly in flocks, and herd mammals graze together in huge num-
bers. The ability to stand crowding is not confined to any one
branch of the animal kingdom, and indeed it varies even among
plants.
In this respect many primates are very primitive. Most if not all
nocturnal species are solitary. Among the diurnal species, most
of the lemurs, if not all, live in groups of from four to fifteen
individuals,7 or more. Apparently all the South American monkeys
are gregarious, and the mean population of howler-monkey troops
has been set at 17.3 monkeys, with a standard deviation of ±6.8
individuals.8 The macaques and open-country baboons live in
groups of about sixty and range from about ten to about two hun-
dred. The larger groups usually include more than one dominant
7 F. Bourliere: Mammals of the World (New York: Alfred A. Knopf; 1955).
8 C- R- Carpenter: “Characteristics of Social Behavior in Non-Human Pri-
mates, TNYA , Ser. II, Vol. 4, No. 8 ( 1942), pp. 251—3.
Adaptation to Crowding ioy
male. Among the apes one species of gibbon, the hoolock, is said
to live in troops; the lar gibbon is monogamous; and the three
great apes live in individual harems. The primates, therefore, are
for the most part gregarious but limited to bands of small num-
bers; and our closest kin, the great apes, live in little kingdoms
of one dominant male and his family and followers.
Although we do not know how our ancestors lived in the days
before speech and tools, it is unlikely, on the basis of comparisons
with primates, that they constituted large bands or troops. The
social unit was probably a small one consisting of one or more
family units. In any case, in the evolution of the human type of
society a threshold must have been crossed when individual fam-
ily bands or households established peaceful relations with other
such bands or households and began exchanging wives.
At this stage each individual came to recognize, know the
names of, and tolerate the presence of several hundred other in-
dividuals. One adult male could then meet another adult male of
a different household, a man of his own age and size, without
challenging him to a fight or creeping away. As many as twenty or
thirty men could get together for a ceremony without the cer-
tainty of serious physical conflict.
The peaceful widening of one’s circle of acquaintances to the
size of a modern, food-gathering breeding unit represented a large
step forward in the process of becoming human. It could not have
been accomplished without language. But even with language, it
was necessary for a healthy and vigorous man to learn to sup-
press his emotions. Not only must he control his speech; he also
had to resist the impulse to ravish an attractive woman or to at-
tack a potential rival. Like language itself, man’s ability to curb
his impulses required genetic changes in the nervous system and
also involved the endocrine system where these emotions are
stimulated. Learning to hunt could have helped a man make this
adjustment, by transferring the target of his aggression from other
males of his group to animals.
Even when human beings had become able to tolerate the
presence of several hundred other individuals, usually in small
groups but occasionally in large ones, adaptation to what zoolo-
gists call crowding did not end. With the invention of agriculture
1Q8 Evolution through Social Adaptation
some peoples came to live in villages and attend intervillage and
tribal markets at which a visitor might see thousands of persons,
many of whom he did not know. In the Bronze Age came cities
and the extreme crowding characteristic of urban communities.
In Asia people who grew wet rice lived in the densely built-up
villages and towns that this miraculously productive cereal is able
to support. The Iron Age brought empires, and the Industrial
Revolution spawned slums. Today vast armies, huge corporations,
and Levittowns channel the interaction patterns of millions of
human beings.
Although human relations have thus been growing more and
more complicated, there has usually been room for two kinds of
individuals. The first and most numerous kind consists of simple
villagers, peasants, petty craftsmen, laborers, and factory workers.
Such people interact with each other in small, face-to-face groups,
informally organized in neighborhoods, work teams, churches,
and clubs. Their patterns of interaction are usually no more com-
plex than those of primitive hunters and gatherers. As civilization
has grown, they have found niches for themselves in its lower
echelons.9
The second kind of individual is the man who has kept pace
with civilization and made it grow. These men rise to high levels
both in hierarchies and in social strata. They are the leaders in
business, politics, religion, education, entertainment, and other
activities. Their circles of acquaintances are national and inter-
national rather than local. Such a man may greet a thousand peo-
ple by name in a single day. He possesses a fine sense of discrimi-
nation and of propriety, knows which words will please and which
will offend, and is able to get others to do what he wants done. He
is capable of choosing reliable lieutenants, delegating authority,
and sleeping soundly after a strenuous day. This man is an
“operator” on a scale impossible in a society like that of the Tiwi.
The same qualities that enable a Tiwi to be successful in ob-
taining women bring him rank, wealth, and fame.
He is in many ways a bright man, but the physical equipment
9 A. F. C. Wallace: “On Being Just Complicated Enough,” PNAS, Vol. 47
No. 4 (1961), pp. 458-64.
Adaptation to Crowding 109
responsible for his success is not limited to the quality of his brain.
Other men as bright as he are failures. His vast energy and his
ability to control it without breakdown depend also on his endo-
crine system. This we know principally by analogy, because of
extensive experimental work done with other mammals.1
In rodents, rabbits, and hares, as well as in other animals, social
pressure — an amount of interaction greater than the animal can
tolerate in comfort — stimulates the part of the brain known as the
hypothalamus, and this organ sends information to the anterior
lobe of the pituitary, commonly called the master gland. The
pituitary in turn reduces its secretion of growth hormone and of
gonadotropins, the substances that stimulate the sex glands to
ovulate, to secrete sperm, and to produce sex steroids, and it also
deforms sperm cells. At the same time the pituitary overstimu-
lates the adrenal cortex. The end product of this neuroendocri-
nological chain of events is stunting, independently of nutri-
tion; reduced fertility; reduced lactation; an altered sex ratio
at birth; increases in susceptibility to diseases; and a higher rate
of mortality.
When animals die as a result of this sequence of events, it is
usually through a rise in the cholesterol level accompanied by
atherosclerosis. This has been shown by Ratcliff e in his autopsy
studies of animals that died during the 1950’s in the Philadelphia
zoo despite an ideal diet high in protein and an attempt to give
each animal as much privacy as possible.2 Still, excess interaction
takes its toll. Individual animals see one another but can neither
attack, flee, nor drive one another away. Thousands of leering and
jeering schoolchildren bait them. Like the overcrowded wild
animals studied by Christian and others, they succumb to endo-
crine disfunction and high cholesterol level, and die young.
High cholesterol level, hypertension, and atherosclerosis are
1 J. J. Christian: Endocrine Adaptive Mechanisms and the Physiologic Regula-
tion of Population Growth, NMRI, No. 60-2, i960.
Christian: “Phenomena Associated with Population Density,” PNAS, Vol. 47,
No. 4 (1961), pp. 428-49.
2 H. L. Ratcliffe and M. T. I. Cronin: “Changing Frequency of Arteriosclerosis
in Mammals and Birds at the Philadelphia Zoological Garden,” Circulation, Vol. 18,
No. 1 (1958), pp. 41-52.
no Evolution through Social Adaptation
prevalent in our own civilization. It has been shown that the
amount of cholesterol in a person’s bloodstream depends less on
fat intake or obesity than on “other biological factors” 3 and that
a tendency to the related diseases listed above is probably he-
reditary.4 High blood pressure is particularly frequent in ur-
banized Negroes both in America and in Africa.5 6 In terms of
animal behavior, all this evidence — much of it quite new and not
yet fully digested by the medical profession — seems to indicate
that individuals vary widely in their inherited ability to resist the
evil effects of large amounts of interaction, and that a higher ratio
of individuals who can withstand it has arisen, by natural selec-
tion, in some populations than in others.0 These differences are not
racial per se, but some races have been exposed to more of this
kind of pressure than others.
In our monogamous society the day has passed when a man of
outstanding administrative ability, whose threshold of tolerance
for crowding is high enough to keep him alive and healthy, can
beget a large number of children. In our society natural selection
seems to work in the other direction, by pruning off those who
cannot tolerate stress. As time, television, and automation move
on, the number of persons who can live as common laborers
dwindles. Every plumber’s son dreams of college and many get
there. Stress is moving down the social scale. As Henry has
shown,7 our mental hospitals fill up faster than our maternity
wards, and in mental institutions reproduction is discouraged.
3 C. B. Thomas and S. M. Gam: “Degree of Obesity and Serum Cholesterol
Level,” Science, Vol. 131, No. 3392 (i960), p. 42.
4M. Kaplan: “Physician Links Hypertension to Inborn Factors, Not Stress,”
NYT, January 27, i960. Kaplan refers to a report by Dr. G. A. Perera at the New
York Heart Association’s annual conference on January 26, i960.
5 R- K. Plumb: “Blood Pressures of Negroes Studied,” NYT, June 3, i960.
Plumb refers to a conference sponsored by the New York Academy of Science on
June 2, i960, in which several papers were read on studies made in South Africa,
Liberia, and the United States but in which no general agreement was reached.
The interpretation is mine.
6 Following a mutation in a single set of gene alleles, the number of fruitflies
that can live together in the same space and with the same amount of food was
trebled in three generations. See H. L. Carson: “Increase in Fitness in Experi-
mental Populations Resulting from Heterosis,” PNAS, Vol. 44, No. 11 (1958),
pp. 1136-41.
7 Jules Henry: “Culture, Personality, and Evolution,” AA, Vol. 61, No. 2 ( 1959),
pp. 221-6.
Ill
Adaptation to Crowding
Many of the inmates of these instutitions could probably have
been adequately adjusted in simpler societies.8
The studies of stress and crowding which have just been re-
viewed are of theoretical value both in general biology and in
human evolution. They point to a physiological mechanism by
means of which animal populations automatically keep their nu-
merical levels constant, even without the aid of predators, which
of course also help. As the selection favors individuals who have
both a stress tolerance and superior intelligence (the two factors
are not otherwise related ) , this may be a mechanism for a general
increase in intelligence in competing animal species.
This non-Malthusian concept adds a new dimension — neuro-
endocrinological competition within a population — to Darwin’s
concept of natural selection. It is selection from within in addi-
tion to selection from without, and in that sense especially con-
cerns general adaptation and evolution by succession, whereas
natural selection for fitness to the environment outside the popula-
tion more closely concerns evolution by branching. This seems to
be a new idea and it will no doubt be challenged, repudiated,
and then widely accepted (I believe) within a few years of the
publication of this book.
As far as man is concerned, this theory helps us understand how
Homo erectus evolved from whatever he was before, and how he
further evolved into Homo sapiens. We now have an idea of what
the other factors” are, beside nutrition and disease, which cause
primitive, marginal populations to drop off in numbers when
faced with invaders and colonists of more elaborate cultures, and
to become extinct, in some cases, through absorption.
The association between tolerance for crowding and high in-
telligence, which, as stated above, is apparently coincidental,
must not be overworked. Neither a businessman who can tolerate
the body heat and noise of a large cocktail party held in a small
room nor a sergeant who can click his heels and transmit orders
in a highly disciplined army is necessarily brighter than a creative
8 The concept that selection for crowding has played a role in culture change
had been thoroughly explored by Schwidetzky before the animal evidence on
which this section is based had become available. See Ilse Schwidetzky: Grund-
> ziige der Volkerbiologie (Stuttgart: F. Enke Verlag; 1950), and also a review of
the same by Paul Leser in AJPA, Vol. 10, No. x ( 1952), pp. 141-4.
112 Evolution through Social Adaptation
scientist who cannot stomach any kind of regimentation and who
prefers to work alone in his laboratory or out of doors. It takes
all kinds to make a world, and it took several kinds of personalities
to make Homo sapiens and to bring him to his present position in
the animal kingdom.
Dwarfing as a Solution to the Problem of Crowding
One of the most controversial subjects in human taxonomy
is the classification of the Pygmies, including principally those of
Africa, the Andaman Islanders, the Semang of the Malay Pen-
insula, and the Philippine Negritos. As yet we have no fully veri-
fied Pleistocene Negrito skeletons to tie us to facts,9 so speculation
has been untrammeled. Some authors give these little folk sepa-
rate subspecific status, but others include them in a larger Negroid
group among the Melanesians, Papuans, Tasmanians, and African
Negroes — in other words, everyone with black skin and curly hair.
To base global relationships on skin and hair alone, without
paleontological support, is dangerous.
The exponents of separate status and, by the same token, a sin-
gle origin as dwarfs, have to postulate early, extensive, and com-
pletely undocumented migrations from Africa to southeast Asia
and Oceania, or vice versa. Those who consider them shrunken
Negroes explain their distribution much more easily, as the result
of independent and parallel acts of dwarfing. No one today, as far
as I know, holds that the Pygmies simply retain the original hu-
man size, under 150 cm. or five feet, whereas the rest of mankind
has grown larger.
As the first two explanations both involve a size reduction from
larger ancestors, it behooves us to study dwarfing in other forms of
life. Dwarfing is common in both plants and animals. Among wild
mammals there are, or have been, dwarf deer in Cuba and Japan;
dwarf elephants in the Philippines, Celebes, and Malta; a dwarf
9 Two Dutch anthropologists found six or more fossil skeletons of small people
in a cave in the island of Flores, Indonesia, in 1955. A report of the tentative
identification of the skeletons as Negritos, and an undocumented estimate of their
age as 30,000 to 40,000 years, has been published only in the press. See Science
Digest, October i960, pp. 62-3; also The Interamerican, Vol. 7, No. 8, Novem- •
ber i960.
Dwarfing as a Solution to the Problem of Crowding 113
mammoth on Santa Rosa Island, California; a dwarf fox on Cata-
lina and Santa Rosa; a dwarf hippopotamus in Liberia; a dwarf
bush-baby (loris) in Africa; a dwarf marmoset in South America;
and a dwarf chimpanzee in the Belgian Congo. The last three,
like Homo sapiens, are primates. Among domestic animals there
are dwarf horses in the Shetlands, Iceland, Oland Island ( Swe-
den), the isles off Rrittany, Sardinia, Corsica, Sable Island, Cape
Verde Islands, Timor, Bali, Sumba, and Japan; dwarf buffalo on
Mindoro and Celebes; dwarf goats on Guadalupe Island, Mexico;
and a galaxy of dwarf horses, cattle, and goats on the Ryukyu Is-
lands.1
Zoogeographically speaking, all these pygmy animals live either
on islands or in small enclaves of tropical forest where their ref-
uges are surrounded by zones of drier terrain. Some of the islands
they inhabit are cool and all are damp; and dampness ensures an
abundance of vegetation. Pygmy men are found only in tropical
forests, which are islands of dense foliage in seas of grass. In
such forests the dense vegetation, fallen and rotting logs, and a
network of hanging lianas make travel difficult; it is easiest for a
small man to move about. Furthermore, little food is available on
the ground. Birds nests, monkeys, fruits, and honeycombs are
high up in trees, and a small man can climb better than a large
man.
In order to continue to exist, a breeding population must re-
main at a greater than critical number. The mammoths of Santa
Rosa Island, for example, which were confined within about 60
square miles, were more likely to maintain the required popula-
tion quota as dwarfs than if they had been full sized. Human be-
ings must attain a minimum population size within a geographical
area, and in addition a minimum number of individuals is needed
if they are to live together in a self-supporting band.
Other special factors, such as the availability of nutritive ele-
ments 2 and the superiority of a small body over a large one in
maintaining thermal equilibrium in areas where a combination of
high atmospheric humidity and lack of wind render sweating
1 A. H. Smith: “The Culture of Kabira, Southern Ryukyu Islands,” PAPS,
Vol. 104, No. 2 (i960), pp. 134—71.
2J. R. de la H. Marret: Race, Sex, and Environment (London: Hutchinson
& Co.; 1936).
ii4 Evolution through Social Adaptation
nearly useless as a cooling mechanism, may add to the advantages
Pygmies share with other dwarfed animals in the tropical forest.
At any rate, their small size must be advantageous or they would
not have become entirely dwarfed, with a 100 per cent fre-
quency of whatever gene controls their stature.
In plants, dwarfing is caused by a single gene mutation, dem-
onstrated in peas, beans, and maize. In peas and beans and in
four of six dwarf strains of maize, the mutation simply reduced
the organism’s capacity to metabolize gibberellic acid. When this
substance was fed to the plants in large doses, they grew to nor-
mal size. The fact that only four of six dwarf maizes responded to
this treatment indicates that dwarfing may be due to any one of a
number of different genes.3
Comparable experiments have been carried on in dwarf strains
of mammals. Several teams of research biologists have shown,
mostly by transplants, that dwarf mice owe their failure to grow to
full adult size to the absence, from the anterior lobe of the pitui-
tary, of either of two kinds of cells (eosinophil and acidophil)
and to a deficiency of growth hormone, and that each condition is
controlled by a single, non-sex-linked gene.4
These dwarf mice were ateliotic ; that is, they were normal for
their species and race in body proportions, with due allowance for
differences of an allometric nature ( see Chapter r, p. 25 ) . Ateliotic
dwarfs occur in many species, including man, and are often infan-
tile in some respects, including sexually. In an experiment per-
formed on human beings, eighteen Caucasoid ateliotic dwarfs,
belonging to a group of related families, grew out of the dwarf
class when, like the mice, they were given pituitary growth hor-
mone.5
Another class of dwarfs is called achondroplastic. Like bull-
3 B. O. Phinney: “Growth Response of Single Dwarf Mutants in Maize to
Gibberellic Acid,” PNAS, Vol. 42 ( 1956), pp. 185-9.
4 R. L. Carsner and E. G. Rennels: “Primary Site of Gene Action in Anterior
Pituitary,” Science, Vol. 131, No. 3403 (i960), p. 829.
P. E. Smith and E. C. MacDowell: “An Hereditary Anterior-Pituitary De-
ficiency in the Mouse,” AR, Vol. 46, No. 3 ( 1930), pp. 249-57.
G. R. de Beer and H. Griineberg: “A Note on Pituitary Dwarfism in the
Mouse,” JG, Vol. 39, No. 2 ( 1940), pp. 297-300.
5 R. L. Schaefer and F. Strickroot: “Endocrine Dwarfism,” Fourth Report,
Endocrinology, Vol. 26, No. 4 ( 1940), pp. 599-604.
The Endocrines and Temperament 115
dogs, they have large heads and bodies, pushed-in faces, and
short, deformed arms and legs with distorted hands and feet. Un-
like the ateliotic dwarfs, they have enlarged pituitaries. The de-
formity of the limbs is usually inherited as a simple dominant,
whereas the facial deformity is inherited in a more complex fash-
ion. Achondroplasia, as this trait is called, occurs in many spe-
cies of animals and something like it is even seen in certain dwarf
trees. In man, as in dogs, the achondroplastic dwarf is sexually
normal.
Human Pygmies, in Africa and elsewhere, exhibit various kinds
of dwarfing. Some are predominantly ateliotic, although sexu-
ally competent, and others show certain achondroplastic features.
The fact that different Pygmy populations vary in these respects
may be added to other kinds of evidence 7 to indicate that the
Pygmies of the world have arisen from separate full-sized ances-
tors in several regions through parallel gene mutations. They are
neither a subspecies nor a single race, and if they are mutually re-
lated it is through their separate full-sized ancestors.
T he Endocrines and T emperament
As everyone who has bred or even worked with dogs
knows, different breeds vary greatly in temperament. A terrier
behaves differently from a bulldog, and setters and retrievers have
special behavior patterns of their own. We know by experience
that these specific breed temperaments are inherited, because the
breeds were selected on that basis. Elaborate experiments have
shown that learning has little to do with them, except insofar as
capacities to learn certain aspects of behavior are inherited.8
Furthermore, and this is particularly pertinent at this point, dif-
ferences in temperament between breeds are accompanied by
diffeiences in the size, form, and histological structure of the
6R. R. Gates: Human Heredity (New York: The Macmillan Co.; 1946), pp.
1320-2. rr
' Gates: “The Melanesian Dwarf Tribe of Aiome, New Guinea,” AGMG, Vol.
10, No. 3 (1961), pp. 277-311. Gates has shown that in both New Guinea and
Africa Pygmies have blood groups similar to those of their full-sized neighbors.
8E. Caspari: “Genetic Basis of Behavior,” in A. Roe and G. G. Simpson: Be-
havior and Evolution (New Haven: Yale University Press; 1958), pp. 103-27.
ii 6 Evolution through Social Adaptation
endocrines, particularly the pituitary, thyroid, parathyroids, and
adrenals.9
Human beings also vary in temperament. It is a common ob-
servation among anthropologists who have worked in many parts
of the world in intimate contact with people of different races that
racial differences in temperament also exist and can be predicted.
Races also differ in the size and weight of endocrine glands, and
in the substances carried in the urine.1 The study of these varia-
tions has just begun, and many readers who believe in the current
dogma that all behavioral differences are due to man’s unique ca-
pacity for learning will find this unpalatable, but the burden of
proof is on them. If such differences are not related to the endo-
crine system, then man is indeed a unique animal.
Parallels Between Animal Domestication
and Social Adaptation
Darwin was deeply interested in the biological results of do-
mestication, and others have carried on a detailed study of this
subject, particularly because of its high economic value. For pres-
ent purposes, one may ask, what has domestication to do with
man? Man is a self-governing animal. But whether an animal is
free or captive, certain modifications take place in its anatomy
9 C. R. Stockard: The Physical Basis of Personality (New York: W. W. Nor-
ton & Co.; 1931).
; The Genetic and Endocrinic Basis for Differences in Form and Be-
havior (Philadelphia: Wistar Institute of Anatomy and Biology; 1941).
D. G. Freedman: “Constitutional and Environmental Interactions in Rearing
of Four Breeds of Dog,” Science, Vol. 127, No. 3298 (1958), pp. 585-6.
H. Oboussier: “Das Verhalten der Hyophyse bei Reciproken von Hunden
gegensatzlicher Wuchsform,” ZA, Vol. 155, No. 5/6 (1955), pp. 101-11.
1 For differences in the size of endocrine glands, see:
E. Loth: Anthropologie des Parties Molles (Warsaw and Paris: Masson et Cie;
i93i)-
W. Freeman: “The Weight of the Endocrine Glands,” etc., HB, Vol. 6, No. 4
(1934), pp- 489-523-
R. Pearl, M. Gooch, J. R. Miner, and W. Freeman: “Studies on Constitution,
IV, Endocrine Organ Weights and Somatological Habitus Types,” HB, Vol. 8,
No. 2 ( 1936), pp. 92-125.
For biochemical differences, see H. E. Sutton and P. J. Clark: “A Biochemical
Study of Chinese and Caucasoids,” AJPA, Vol. 13, No. 1 (1955), pp. 53-65.
These references are only a small sample of the pertinent literature.
Animal Domestication and Social Adaptation 117
and physiology as a result of cultural protection. Klatt,2 3 * * among
others, has shown that animals in a state of domestication respond
to crowding, and to a reduction in mobility and in sensory percep-
tion. They do not need to walk far for their food, and some of it
is even brought to them; nor do they need to watch for predatory
carnivores, because men and dogs protect them. They do not have
to seek shelter in bad weather, because they are driven into barns
or caves.
These factors bring about a reduction of bone density, an in-
crease in adipose tissue, a reduction in the length of the snout and
in some cases in tooth size, and a reduction in brain size of 10 per
cent, 20 per cent, or even 30 per cent of the measurements for
wild foims. This has been established in the case of dogs, ferrets,
pigs, ducks, and cats. When house cats become feral, larger brains
reappear. Domestic animals do not ordinarily use their senses
as much as their wild counterparts do. This is indicated by reduc-
tions in the area striata of the brain, which is concerned with vi-
sion, and by smaller eye sockets (orbits), and smaller auditory
apertures on the skull. In the dog, which is a working animal ( or
used to be), the brain is smaller than the wolfs but shorter and
higher, and the forebrain is larger. Klatt interprets the growth of
the canine forebrain to a selection for the capacity to do the work
hunters and shepherds require of him.
In our study of fossil skeletal material it will be interesting to
watch for possible changes in man comparable to those seen in
domestic animals. We shall find long skulls, protruding occiputs
( area striata), heavy brow ridges, large orbits (eye sockets), and
large auditory canals in the earliest forms, which existed when
2 Berthold Klatt: Haustier und Mensch (Hamburg: Richard Hermes Verlae-
1948). 6’
3 The reversion of domestic animals to wild prototypes once they have become
feral can be demonstrated by experiments on insects, particularly on body lice.
The head louse, a human parasite, is so different morphologically from the body
louse that they have been considered different species ( Pediculus capitis and
P. vestimenti) . Yet, when head lice are forced to live on the body they change
form drastically over several generations, and when they move back to the scalp
they gradually resume their original form. The difference in environment between
a covered and an uncovered part of the body may be compared to wild and do-
mestic conditions. H. Levene and T. Dobzhansky: “Possible Genetic Difference
between the Head Louse and the Body Louse ( Pediculus humanus L.) ” AN
Vol. 93, No. 873 (1959), pp. 347-53-
1 1 8 Evolution through Social Adaptation
men were living close to nature and few could sit in the camp and
be fed by others. But we cannot expect to see many of the changes
which accompany domestication appear in males very early in the
Pleistocene. However, as women are protected more than men,
these changes could appear earlier in the female than in the male
skeletons. Later on, with the advent of villages and cities and pre-
cooked frozen meals, these modifications may appear in numbers.
At any rate, this is another possible physical result of social adap-
tation in man to add to an initial increase in brain size, subse-
quently reduced in some populations; to an adaptation to crowd-
ing; and to dwarfing.
The Unique Adaptations of the Genus Homo
Like other animals, Homo has adapted himself to
living in his terrestrial environment and with other animals of
his own species. Many of the forms these adaptations have taken
follow common garden rules of biology, including both zoology
and botany; probably all of them do. What is unique is the way
man has interrelated them, or more accurately, the way the forces
of nature have interrelated them for him.
He has stretched the capacity of a hairless tropical primate to
its physiological limits in conquering unfavorable climates. He has
learned how to make fire and shelter, carry water, and store food.
He has learned to talk and to think in terms of language, and to
live in larger communities than any of his fellow primates. He has
found ways to tolerate crowding and a certain amount of regi-
mentation. All these are cultural in a sense but in another and
more inclusive sense they are biological because they have af-
fected the genetic structure of the human organism. Culture and
biology are parts of a single picture and both require equal treat-
ment in the review of human racial history that follows.
THE ORDER OF PRIMATES
Primate Studies and the Classification
__ _ of Human Races
_l_ he purpose of this book, which must not be lost from
view in the dense foliage of information — drawn from many dis-
ciplines— needed for its composition, is to trace the descent of the
living races of Homo sapiens. To discover the genetic relationships
among these living races, we shall attempt to follow them back in
time to ancestral fossil races, derived in turn from one or more
prototypes in the extinct species Homo erectus, itself descended,
through the mists of the fossil record, from still earlier prehomi-
nid forms. This must be done because it is possible that the an-
cestors of the living races parted company very early in the his-
tory of our genus.
We must also learn certain specific facts about the other living
members of our order, prosimians, monkeys, and apes, because the
primates as a whole are variable in anatomy, physiology, and
behavior. Some of these variations are mentioned briefly in the
preceding chapter. A more detailed study of them may disclose
pertinent data about the descent of our genus and also about the
possibilities within the total primate gene pool for all kinds of phe-
nomena, such as the lack of a tail, different ways of walking erect
(when this is done occasionally), the capacity to live in groups,
communication, different amounts of skin pigment, a prominent
external nose, and so on. These variations recur in some races of
man but not in others, and they may be primitive, neotenous, spe-
cialized, or adaptive. We shall be in a better position to under-
stand these traits in man if we first examine them in other pri-
120
The Order of Primates
mates. Such a preliminary study will also enable us to judge
between close kinship and parallelism with greater confidence.
The Classification of Primates
If the relationships among the units in any set of phenomena
are to be studied, these units must be classified. Had chemists not
classified atoms and molecules, we would have no modern in-
dustry. If zoologists and paleontologists were not able to classify
the order of primates, we could not hope to formulate a valid hu-
man taxonomy. Unfortunately for present purposes, no ironclad
classification has yet been made; but the problem is not as great
as it seems, for most of the difficulties involve the designation of
the higher categories: suborders, infraorders, superfamilies, and
families, which interest us least. There is little disagreement about
genera and species, which are most pertinent to our study.
The classification given in Table l is based on Simpson’s ( 1945),
with emendations from other sources, chiefly Fiedler (1956).1
When Fiedler and Simpson are in disagreement, this fact is indi-
cated by initials in parentheses: (F) and (S).
The Prosimians
All modern taxonomists group the lemurs, lorises, and
tarsiers, with or without the tree shrews, in a convenient category,
that of the prosimians, which distinguishes these lowly members
of the primate order from New World monkeys, Old World mon-
keys, apes, and men. They are of particular interest because they
1 From 1945 on, the principal sources on primate taxonomy are as follows.
G. G. Simpson: “The Principles of Classification and a Classification of Mam-
mals,” BAMN, Vol. 85, 1945.
W. L. Straus: “The Riddle of Man’s Ancestry,” QRB, Vol. 24, No. 3 (1949),
pp. 200-23.
W. C. O. Hill: Primates, Vol. I, Strepsirhini (Edinburgh: University Press;
1953) and Vol. II, Haplorhini (Edinburgh: University Press; 1955).
W. Fiedler: “Ubersicht fiber das System der Primates,” in D. Starck and S.
Karger, eds.: Primatologia, Vol. I (Basel: S. Karger; 1956), pp. 1-266.
My scheme will of course be superseded when Simpson’s revision of his 1945
work is published.
The Tree Shrews
121
are exceedingly primitive mammals, in many ways not far ad-
vanced beyond the marsupials, and because they have much in
common with our closest mammalian kin, the insectivores. The
ancestors of these prosimians formed the base from which the
higher primates developed. Knowing about them will help us un-
derstand the relationships among the more advanced forms.
The Tree Shrews
Standing near the taxonomic frontier between the pri-
mates and the insectivores are the tree shrews, which have a su-
perfamily and family of their own, divided into two subfamilies
and four genera. The tree shrew proper, genus Tupaia, has eight
species; the smooth-tailed tree shrew ( Dendrogale ) has two; and
the Philippine tree shrew ( Urogale ) and the pen-tailed tree shrew
( Ptilocerus ) have one each. All are confined to the Oriental re-
gion, principally to southeast Asia and Indonesia.
Being among the smallest of mammals, they have a high meta-
bolic rate and require much food. If two males are left together in
a confined space overnight, one will eat the other. They are enor-
mously energetic, voracious, belligerent, amorous, irascible, and
omnivorous, and particularly partial to a diet of insects, which be-
fits their size. Their excesses of courage, wrath, and libido consti-
tute a caricature of uninhibited human behavior unparalleled by
any larger and fully accredited primate.
Like the insectivores they have claws on all ten toes; other
primates have at least some nails. Their tooth formula is — 1-3:3,
3:i.'3:3
which is zoological shorthand for: upper incisors = 2, upper ca-
nines = 1, upper premolars = 3, upper molars = 3; lower inci-
sors = 3, lower canines = 1, lower premolars = 3, lower molars
= 3- Thus, each of the four kinds of teeth commonly found in
mammals is enumerated for one side of the face. To calculate
the total number of teeth one would multiply by 2. The tree
shrews, then, have 38 permanent teeth. We have 32. The tree
shrews uniquely, among primates, have three lower incisors on
each side. All the others have two or one.
MAP 4
living prim ate s
MACAQUES -i
CA
M langurs * IY ✓
S 1
^ r
CC3»
~~ctj
/ilV
— r . c\
/ Hill
BABOONS
;■;' ^.mountain /
mwmwmm.QOKiLiAS _
P«GA1V Q>
^chimpanzee 4 f /
) 4 /4
%-Ltarsius
n*&v /.i .
YiGIBBON
. . I jnEit.v'i
TUPAIA \fW /l^/
(TREE /S PROBOSCIS/
/If
^lIP
SI AM A NO i
Lemurs. t;.::-;-;-,;-:-l A/'w Monkeys.
Ethiopian Fauna-- Lorises, Galagos, Colob us Monkeys,
Cercopithecus Monkeys , "Baboons, Chimpanzees, Gorillas .
Oriental Fauna-. Tree Shrews, Lorises, Tarsiers, Langurs,
Macaques (including Barbary Apes), Gibbons ,siamangs,
and Orangutans.
126
The Order of Primates
The male tree shrew has a large ( for him ) and pendulous penis,
with external testicles. The female has a double-chambered uterus
and each placenta has two discs. The tree shrews proper have
three pairs of nipples, the smooth-tailed ones two pairs, and the
Philippine and pen-tailed ones a single pair each. Their eyes are
large and set at right angles to each other; the binocular vision of
the higher primates is missing. Yet the visual area of the brain
( area striata) is large compared to that of the insectivores, and
the olfactory region is relatively small. In some species the exter-
nal ear is rounded as in man. All in all, these little animals seem to
be a curious mixture of features left behind and features to come.
The Lemurs
The superfamily of Lemuriformes includes three fami-
lies, all of which live exclusively in Madagascar. In the shelter of
this secluded island, where no other form of mammal gave them
serious competition, the lemurs (using the word in its inclusive
sense to cover all three families) specialized, just as the marsu-
pials did in Australia. As with other island faunas, many of the
larger species disappeared when the island was opened to conti-
nental animals by the arrival of man.
Most of the lemurs are quadrupedal climbers with small brains
for primates, a good sense of smell, a tacked-down upper lip and a
minimum of facial expression, a wet nose, tactile whiskers, non-
stereoscopic vision like that of the tree shrews, and hands and feet
in which the second and third fingers and toes are clawed but the
other three digits end in nails like those of the other primates. In
some species the second digit is rudimentary; and in most the
longest is the fourth, instead of the third as in other primates and
in most five-toed mammals.
The long, narrow, V-shaped lower jaws of most lemurs contain
one tooth less on each side than those of the tree shrews and one
more than those of Old World monkeys, apes, and men. Their
dental formula is
2: 1 :3:3
2: 1:3:3
In the lower jaw the four incisors and
two canines point forward in unison to form a stabbing or shear-
The Lemurs
127
ing device known as a dental comb, and the first lower premolar
has become enlarged to assume the duties of the lower canine in
other animals. These are very aberrant specializations.
However, one family, the Daubentonia, with but a single
species, the aye-aye, has an even more specialized and greatly
reduced rodentlike dentition, with this formula: —
1
18 permanent teeth as compared with 36 for the other lemurs and
32 for men. Its incisor teeth, which are huge, grow with wear,
like those of rodents. With them these animals cut through the
hard stalks of canes, especially sugar cane, to get at the sweet
: 1 :3
:o:o:3
, totaling
Fig. 3 The Skull of An Aye-
Aye. The Aye-Aye ( Daubento-
nia ) is a highly specialized lemur
of Madagascar which has chisel-
like incisor teeth. It uses them
like a rodent. ( Drawing after
Flower and Lydekker, i8gi.)
sap. They also use the incisors in chiseling wood, like wood-
peckers, to uncover grubs, which they pull out with long, slender,
highly specialized middle fingers. Except for the great toe, which
bears a nail, all their digits end in claws.
Unlike most primates lemurs have a breeding season, and one of
them, the mouse lemur (Cheirogaleus) , sleeps through the hot
season, or more technically, it estivates. The uterus of the lemur,
like that of the tree shrew, is forked. Whereas all the other pri-
mates, including the tree shrew, have disk-shaped placentas, the
lemurs have bell-shaped ones. Also uniquely among primates, the
lemur’s placenta fails to emerge with the infant at birth. Twins
are frequent, and some genera have four nipples each, others only
two.
The lemurs are too specialized to have much bearing on pri-
128
The Order of Primates
mate origins, but they serve as classic illustrations of several prin-
ciples. Ranging in size from the stature of a mouse to that of a
man, they probably failed to increase much in intelligence once
they had reached Madagascar. Because of lack of competition
with other terrestrial mammals, intelligence was not at a pre-
mium. The lemurs achieved as complex a pattern of evolution and
as great a proliferation of species as could be expected on an is-
land the size of Madagascar. The two sets of organs most con-
cerned with locomotion and feeding became the most specialized :
the hands and feet, and the teeth. The lemurs were no match for
man and the other invaders that accompanied him from the East
Indies and Africa. The largest species were the most vulner-
able and the first to disappear.
The Lorises
Much more widely distributed, and in a sense less special-
ized, than the lemurs proper are the members of the superfamily
of Lorisiformes. The family of Lorisidae has four genera, and that
of Galagidae but one. The Lorisidae, or lorises proper, include the
slow and slender lorises of southeast Asia and India and the
angwantibo and potto of West Africa. These are all small animals,
squirrel- to cat-sized, heavily furred, with round heads and short
ears, tails rudimentary or absent, and nails on all digits except the
second toe. They breed throughout the year. They are all noctur-
nal and solitary, and they are all slow movers, especially the slow
loris and potto. Seeing them creep along a limb is like watching a
slowed-down movie.
Their slowness is due to a lowered metabolism, which is helped
by the rete mirabile ,2 or net of blood vessels in the extremities
which produce a heat exchange between outgoing arterial and in-
coming venous blood, as occurs, in a more simplified form, in the
extremities of Australian aborigines. Like the sloth, another slow
animal, the lorises need their fur, even in the tropics, to keep them
warm under these circumstances.
2 P. F. Scholander: “The Wonderful Net,” SA, Vol. 196, No. 4 (1957), pp.
96-107.
The Tarsiers
129
With large thumbs pointing straight out sidewise and rudimen-
tary index fingers, they creep about by grasping limbs, in search of
leaves, buds, grubs, birds’ eggs, and other immobile or slow-mov-
ing food. I once saw a potto bite clean through the thumbnail of a
man who was holding it and was too busy talking to notice its
snail-like movements. Uniquely among mammals this animal has
four or five long, pointed vertebral spines, which stick through the
skin of its neck and shoulders. By raising them when attacked, it
can inflict a nasty wound on a predator.
One of the Asiatic species, the slender loris ( Loris tardigra-
dus ), which is not quite as slow as the others, parallels man in that
its body length is in ratio to its leg length and that it has no tail.
The galagos lack the lorises’ low metabolic rate. They are quick-
moving animals with large, erect, and mobile ears; they make
their way through the trees by long leaps. Like several other Afri-
can mammals, including man, they come in two sizes, regular and
pygmy. One species, the needle-clawed euoticus ( Galago elegan-
tulus) , has nails like the others, but each nail has a ridge running
down the middle which ends in a sharp, clawlike point. They are
particularly great leapers, and use these needle points for grasp-
ing branches when they land. All galagos have a secondary un-
dertongue. That of euoticus is fringed like a comb; with it the ani-
mal cleans its fur.
The Tarsiers
The superfamily of Tarsiiformes is represented today by
a single genus and what is probably a single species,3 Tarsius spec-
trum. Tarsius is an extremely important animal. In its fossil form it
is believed to have served as an evolutionary bridge between the
Lemuriformes and all the higher primates. Also, according to the
celebrated anatomist Wood Jones, man is descended directly
fiom a tarsioid ancestor without intervening benefit of monkeys or
apes.4
Whatever his position in the progression of primate evolution,
3 Other species have been proposed. See Fiedler: op. cit., pp. 125-7.
4F. Wood Jones: Man’s Place Among the Mammals (New York: Longmans,
Green, & Co.; 1929).
130
The Order of Primates
Tarsius today is a pocket-sized prosimian equipped with a pair of
huge, almost but not quite stereoscopic eyes; hands and feet lined
with spongy pads for clinging to limbs; and a specialized length-
ening of the tarsal bones, whence his name. These are bones of
the ankle, heel, and the back part of the arch of the foot. In man,
compared to other mammals, they are usually short; but in Tar-
sius they are greatly elongated to facilitate hopping, at which he
is a champion. He retains claws on his second and third toes: the
transition from claws to nails in his suborder is not complete.
An animal that hops in the dark needs better vision than a
night crawler, which can guide itself partly by touch. Hence the
huge eyes of the tarsier. To go with these, the visual cortex of his
brain is correspondingly enlarged, and the back of his skull
rounded, as in man. Another humanlike feature is the reduction
of his jaws in relation to the size of his head. Tarsius crushes grass-
hoppers with his fingers, puts them in his mouth, and swallows
them, just as people swallow premasticated food. His teeth are
somewhat reduced in number; their formula is — — ■ The
1: 1:3:3
upper median incisors are wider than the upper laterals, as in
man, but the small jaws converge in a V-forrn in a very unman-
like fashion.
Clutching the shaft of a twig large enough to hold his inconse-
quential weight, Tarsius sits erect, looking about him for even
tinier prey. He can turn his head around i8o° and gaze directly
behind him, thus completely covering the field. In order to rotate
to this extent, his head is hafted in the center, and his foramen
magnum is therefore placed forward of its location in other pri-
mates except man.
The tactile whiskers of the lemur remain, but the wet nose is
gone, along with the long frenum tacking down the upper lip.
Whereas in the lemur and the loris the bony ear has two pieces,
the bulla and annulus, in Tarsius these are fused to make an ex-
ternal auditory meatus, as in man. All in all, Tarsius anticipates
the higher primates in several features, some perhaps in a truly
evolutionary sense, others independently developed by each, in
parallel fashion, because of a special way of life.
The Living Platyrrhines: the South American Monkeys 131
The Living Platyrrhines: the South American Monkeys
The platyrrhines evolved in a tropical paradise much
larger than Madagascar and nearly as free from competition.
They produced a variety of primate forms based on a prosimian
pattern, mimicking, in locomotion particularly, the monkeys and
apes of the Old World tropics. Because their nostrils, like those
of lemurs and many other mammals, are widely separated by a
broad septum, they are called platyrrhines, in contrast to the Old
World monkeys, or catarrhines, whose nostrils are pinched to-
gether, as, to a certain extent, is true in man. Their basic dental
formula is also lemuroid: — — — . Their placentas, on the other
2: 1 13:3
hand, are single deciduous discs, as they are in Tarsius.
The most primitive members of the platyrrhine suborder are
the marmosets, the three genera of the family Callithricidae.
They are small monkeys with lemurlike body proportions, and a
tail half as long again as head and body combined. The male, who
has a scent gland at the base of his tail, raises it as a signal of
sexual interest. Uniquely among primates, the female habitually
gives birth to two or three young at a time and if three are born
the mother kills one. Except at feeding time the father carries the
babies, and when they are ready for weaning he chews food for
them.5
The marmoset has claws on all digits but a great toe and a
thumb with feeble powers of adduction. The face still has tactile
whiskers, and the whole facial musculature consists of a single
subcutaneous sheet, the platysma, which, as in lemurs and other
primitive mammals, is not differentiated into separate muscle
bundles as it is in higher primates, including man, who communi-
cate partly by facial expression. The marmoset’s dental formula is
reduced, with only two molars on each side of each jaw. Like the
galagos, the marmosets have a pygmy form.
The rest of the South American monkeys, the Cebidae, have
5 W. E. Edwards: Why the Marmoset Grew Her Tail, paper read at Am. Assn.
Phys. Anth. meeting May 13, 1960.
132
The Order of Primates
2 ; 1 ; o • O
the standard prosimian dental formula, — — — ' , and nails on all
2 : l : 3 : 3
fingers and toes. The thumb is either nonopposable or absent.
The eyes of the marmosets, like those of the prosimians, point a
little to one side. The eyes of the cebid monkeys, however, point
forward to produce a single stereoscopic image, as in the ca-
tarrhines, including ourselves. Furthermore, in the marmoset eye
the retina is largely undifferentiated.
The retina of the cebid monkeys, on the other hand, has a cen-
tral depression called the fovea which is necessary for fine focus-
ing. This is richly equipped with cones, the kind of nerve ends
used in color vision. As both the platyrrhines and the catarrhines
evolved separately from the prosimians, stereoscopic vision and
color vision were acquired independently in the higher primates
of the Old and New Worlds. Only one genus of cebid monkeys,
Aotus, lacks color vision; this animal is nocturnal and does not
need it.
The cebids also parallel the catarrhines in brain development,
and in intelligence. In both groups progressive series can be laid
out, from large to small brains, and from simple to complex con-
volution patterns. The details of brain structure differ in the two
groups, regardless of size or complexity of cortical folding. This
observation leads to the thought that if brains can evolve inde-
pendently in size and complexity in two related suborders, then
the same kind of parallel evolution could take place in equally iso-
lated populations of smaller taxonomic magnitude, such as gen-
era, species, and even subspecies. This concept is vital for the un-
derstanding of the origins of human races.
In their means of locomotion the cebids have diversified widely.
One of their adaptations is unique; the others parallel those of the
Old World monkeys and apes. The unique feature is the long pre-
hensile tail found in the three most specialized forms, the capu-
chins ( Cebus ), the howler monkeys (Alouatta) , and the spider
monkeys ( Ateles ). They use the tail in locomotion as a fifth limb,
and they can even hang by it. With it the spider monkey can pick
up food and put it in his mouth. Such a tail is particularly useful
in the rainy season, when forests are flooded. The monkeys can-
not go down to the ground and if they fall they may drown.
The Living Cercopithecidae: Old World Monkeys 133
In the use of their limbs the Cebidae fall into two categories,
those that resemble the Old World monkeys and those that ape
the apes. From Callimico to Callicebus to Cebus is a linear evolu-
tionary sequence to be expected more of a fossil series than of a
living assemblage.
One sideline is that of the Pithecinae, which have short tails;
another is that of the Alouattinae, or howlers, which eat leaves
like the Colobinae and vocalize as loudly as gibbons. Their noise-
maker is a chamber in the throat enclosed by a huge hyoid bone
and a pair of swollen and everted gonial corners in the mandible.
Like the apes, the Atelinae brachiate, that is, they move
through the trees by swinging hand over hand between branches.
This mode of locomotion is optional with Lagothrix, the woolly
monkey, whose thumbs are still large and whose forelimbs are
shorter than his hindlimbs, as in nonbrachiating forms. Brachy-
teles, the woolly spider monkey (whose nose is narrow like those
of catarrhines ) , has somewhat longer forelimbs and brachiates
more frequently. Ateles the spider monkey, the largest of all and
at the end of the evolutionary line, is a full-time out-and-out
brachiator except when he is swinging by his tail. He has ac-
quired a gibbonlike body size and body build with a short trunk,
long limbs, and a particularly long forearm. As his hands are used
primarily as swinging hooks, Ateles, who can pick things up with
his tail, has lost his thumb, and when at rest his four fingers are
kept in a hooked position by short tendons that stop the fingers
from straightening out except when the palm is bent forward.
This apelike specialization prevents strain in his finger muscles
when he hangs from his hands for long periods. On the ground he
walks on his mid-phalangeal finger joints, like an ape.
The Living Cercopithecidae: Old World Monkeys
The catarrhines, including the Old World monkeys,
apes, and men, have in common the type of nose for which they
were named. In it the nasal passages are set close together and
parallel, separated only by a narrow septum; and the nostrils, of
variable width, are set fairly close together and point downward.
134
The Order of Primates
The catarrhines have a reduced sense of smell in comparison with
the other primates. They all are diurnal and have stereoscopic
color vision and a fovea. They have a fused bony ear with an ex-
ternal auditory meatus. Another feature of the catarrhines is a
2 I 1 ! 2 I
dental formula, — — - — — , which they alone possess and which
2: 1 12:3
represents a reduction by one premolar ( the first ) from the lemu-
roid ancestral form.
Despite these common features, the Old World monkeys, apes,
and men cannot be included in a single family, or even a super-
family, because of two fundamental differences, (l) The Old
World monkeys have a two-disc placenta, also found in tree
shrews. The apes and men have the usual one-disc form in com-
mon with the tarsiers and platyrrhines, although a two-disc
placenta occurs as a rare anomaly in man. (2) The family pe-
culiarity of the Cercopithecidae is their molar teeth.
All primate molars have cusps, or moundlike projections, on the
occlusal or grinding surfaces. These cusps tend to be worn down
through use during life. Although the number of cusps varies in-
dividually, racially, and otherwise, between three and six, there
are characteristically four principal ones, one in each corner. The
four sides of the tooth bounded by these cusps are known as the
labial, on the tongue side; the buccal, on the cheek side; tire
mesial, or front; and the distal, or back.
Uniquely in the family of Cercopithecidae, the two mesial
cusps are linked by a crossbar into a ridge, known technically as a
loph, and the two distal cusps form a second loph. Such teeth are
called bilophodont. Among the Old World monkeys the first and
second upper and lower molars are characteristically bilophodont,
and the third of either or both jaws may or may not be. The molars
of apes and men lack bilophodontism. As both the two-disc pla-
centa and bilophodontism are specializations, the absence of
these features in apes and men is commonly taken to indicate that
the ancestors of apes and men must have branched off the com-
mon ancestral Old World simian stock before the ancestors of the
Cercopithecidae had acquired these features.
A third peculiarity of the Cercopithecidae which is almost but
not quite exclusive to them is the possession of ischial callosities.
The Leaf-eating Colobinae
135
Fig. 4 The Molars of Old World Monkeys and Apes. Left: A bilophodont
lower molar, typical of the Old World monkeys. The molar has four cusps, in two
lophs or pairs, each loph linked by a connecting ridge. Right: A Dryopithecus-type
lower molar, typical of apes and men. Such a molar has five cusps, situated inde-
pendently around the edge of the crown, with the fifth to the rear.
These are bare, rough spots on the animal’s skin, which cover the
projections of the ischial bones on which it sits down. As the
presence of this built-in seating pad is matched by a roughened
area on the underlying bone, it can be detected in fossil forms.
All the Cercopithecidae have these callosities, but so do other ani-
mal forms. Small callosities are to be seen, although they are
partly concealed by fur, in the gibbon, and they turn up now and
then in chimpanzees. Ischial callosities have nothing but their
general location in common with another and much more con-
spicuous primate feature, sexual skin, which is found in some gen-
era of both monkeys and apes.
The Leaf-eating Colobinae
The Cercopithecidae are subdivided into two sub-
families, the Cercopithecinae and the Colobinae. The latter is
limited in Africa to a single genus extending from West Africa
across the Congo to Ethiopia, but it has three genera in Asia,
where they comprise the majority of all fully arboreal monkeys.
Like the South American howlers the Colobinae specialize in eat-
ing leaves. (The langur and snub-nosed langur also eat fruit, and
the latter steal food from men. ) Their adaptation involves princi-
pally the digestive apparatus. The colobinian stomach is very
large and, when full, contains a load of leaves equal to one third
of the animal’s body weight, or ten times as much food as man re-
quires. Resembling a modern combination living room, dining
room, and kitchen, this stomach is partially divided into semi-
The Order of Primates
1 36
detached subchambers by a series of constrictions. The liver,
crowded by the enormous food container, is lobeless, and is sepa-
rated from the diaphragm, as in the human fetus.
As leaves require chewing rather than biting, the incisors and
canines of these animals are relatively small, and the molars pro-
portionately large. The chewing mechanism, which calls for side-
wise grinding as well as up-and-down motion, involves a stronger
development of the masseter muscles of the face than of the
temporal muscles of the vault. In this whole dental and muscular
complex they resemble the Hominidae, including ourselves. Fur-
thermore, two of the three Asiatic genera of Colobinae have also
acquired prominent external noses, which make them look like ca-
ricatures of people. In Nasalis, the common monkey of Borneo, the
male starts off in childhood with a tip-tilted sensitive-looking little
nose; in early maturity the nose stands out straight and strong,
and it collapses grossly and pendulously to lip level when the ani-
mal is approaching the end of its life cycle. The female’s nose is
much smaller. Rhinopithecus, the snub-nosed langur of Tibet and
China, retains an uptilted nose throughout life.
The stenophagic diet of the Colobinae, which provides them
with an abundance of easily acquired food, thus permitting dense
population growth,6 limits them to forested regions of little sea-
sonal change in which the foliage is nondeciduous. Although
barring much of Africa, this diet permits the Colobinae to extend
their range outside the tropics into the cool Himalayan and
trans-Himalayan forests, which are inhabited by two kinds of lan-
gurs. Like other narrow ecological specializations, this diet is an
evolutionary blind alley which primates of more than one cate-
gory have entered but from which none has emerged by produc-
ing a higher form.
The Cercopithecinae
Just as the forests of the Oriental region are mainly inhabited
by the Colobines, so those of Africa are typically the playground
of arboreal Cercopithecines. This may be partly due to the rela-
tively small seasonal change in rainfall in Asia and Indonesia
6 In Africa hundreds of thousands have been killed for fur.
137
The Cercopithecinae
and to the availability of green leaves there throughout the year.
In most of Africa, on the other hand, alternate wet and dry sea-
sons cause many species of trees to lose their leaves once a year,
so that monkeys which eat fruit, grubs, and other nonleafy foods
have an advantage over leaf eaters.
Beside being omnivorous, the Cercopithecines have several pe-
culiarities, including in most species cheek pouches in which they
carry food, and in all species superorbital notches, that is, a notch
in the top of the bony orbit of each eye which permits the shel-
tered passage of a nerve, a vein, and an artery. In man and the
apes these may pass through a foramen, or small circular opening,
instead. All the Cercopithecinae have well-developed, opposable
thumbs, and many have relatively short tails. Their brain develop-
ment is variable between genera and reaches its greatest height
among primates other than apes and men. They use elaborate
techniques of communication, both by voice and by facial expres-
sion.
The subfamily can be further divided into two groups accord-
ing to habitat and means of locomotion, the arboreal and the ter-
restrial. The arboreal group, which is exclusively African, includes
three genera, Cercocebus, Cercopithecus, and Erythrocebus,
which are divided into many species and races. These monkeys
vary greatly in hair length and thickness, coiffure styles, and
color, with greens, yellows, reds, blacks, whites, and various com-
binations of these variations. The smallest is the talapoin of the
genus Cercocebus, which shows many parallels to the New World
Cebus. It has a globular brain case, a long lumbar section of spine,
little protrusion of the jaws, and small teeth. From this animal a
progression may be found to the patas monkey, Erythrocebus,
a red-haired monkey of East Africa which lives in open country. It
has a protruding snout and a short tail, and is as much at home on
the ground as in the trees. It is thus transitional to the genera
which live primarily on the ground, the macaques ( Macaca ),
baboons ( Papio ) , and gelada ( Theropithecus ) .
These three have become specialized for a way of life not
known among the primates hitherto described — life on the
ground. Although they can and do climb trees without difficulty
— baboons even sleep in trees when these are available — monkeys
The Order of Primates
138
of these genera walk about on all fours, with the palms of their
hands flat on the ground, seeking food in bushes, under stones,
and elsewhere on the surface of the earth. Some of them even dig
shallow-growing roots with their stout fingers. Thus they are able
to occupy successfully lands unavailable to other subhominid pri-
mates, and to live in the forest as well. Many of them seem to pre-
fer rocky terrain to flat country, and they are accomplished rock
climbers. All of them can sit up when resting, and all have re-
tained full mobility of their fingers despite their quadrupedal gait;
they are, in fact, more dexterous than brachiating monkeys and
other treetop climbers including the howler. Such is the construc-
tion of their jaws that, when sitting erect, they can open their
mouths wide only by throwing back their heads. They all have
muzzles and large teeth, some being more doglike in this respect
than others. They are vigorous, belligerent, amorous, and formid-
able fighters, particularly in packs.
The least specialized anatomically and the most widely dis-
tributed is the genus Macaca, which is the second most widely
ranging of all primate genera after Homo. Macaca has many
regional forms. Reading from left to right on the map, the first
is the Barbary ape, native to Gibraltar and parts of North Africa/
particularly the rocky face of Mt. Meggu, behind the city of
Shehshawen in northern Morocco. At Mt. Meggu the Spanish
soldiers used them for target practice during the RifBan War of
the early 1920’s, and these monkeys characteristically carried
away their wounded, probably mostly young ones. The Barbary
ape is a shaggy, yellowish-brown animal with no tail.
The second is Macaca mulatta, formerly called M. rhesus,
which ranges east from Pakistan over much of India. It is the
common laboratory monkey in whose honor a set of blood groups
has been named, and hardly needs description. It inhabits both
open country and forests; and a related form ranges high in the
Himalayas along with the langurs.
A species called Macaca arctoides, large and hairy, lives as far
7 The widespread notion that the Arabs brought this animal to Gibraltar, either
from North Africa or from as far away as Pakistan, has no historical basis and is
refuted by paleontological evidence. See N. C. Tappen: “Problems of Distribution
and Adaptation of the African Monkeys,” CA, Vol. l., No. 2 (i960), pp. 91—120.
The Cercopithecinae 139
north as Szechuan in China, and others even get to the Amur
River and Korea. The macaques of Japan dig for food under the
snow in winter and swim in the sea for shellfish in summer. A
Formosan macaque lives in caves along the coast and is semi-
aquatic, diving for crabs and shellfish and special seaweeds. The
crab-eating macaque of Indonesia and the shores of southeast
Asia, Macaca irus, lives in mangrove swamps, and swims after
sea food like the others. In Celebes lives the so-called Moor ma-
caque ( Macaca Maura), a black species with a stumpy tail. It
goes about in packs and is said to hunt other animals.
In sum, the genus Macaca shows as much range of behavior as
of habitat. It is one of the most versatile of mammals, not only
among the primates but in any order. Mainly tree-borne in south
India and Ceylon, mainly terrestrial in most other places, it is an
alpinist, a swimmer, a hunter, a fisher, a mollusc gatherer, and it
eats the whole menu from shellfish to nuts. No wonder, then, that
medical researchers have favored it as their chief laboratory ani-
mal. If its versatility was a result of living away from the forest
and on the ground, this may help explain the wide distribution of
our own ancestors after they, too, had ceased to be arboreal.
The other terrestrial Cercopithecinae are the black ape, Cyno-
pithecus, and the baboons. The black apes inhabit Celebes, Batch-
ian, and some of the southern Philippines. Although related to the
macaques, they have developed muzzles and cheek swellings that
make them look like baboons. They, too, include sea food in their
diet.
The baboons, of the genus Papio ( P . papio, comatus, cyno-
cephalus, doguera, and hamadryas ), are limited to Africa and
southwest Arabia. They are large animals with tails shorter than
the combined length of their body and head, doglike muzzles
equipped with large teeth, including long, daggerlike canines in
the male. The face is bare, ribbed, and red, and the posterior
adorned with a large, continuous, pillowlike crest of ischial cal-
losities which, as Washburn has observed,8 serves as a cushion on
which the bulk of the animal’s weight rests while it is sleeping in
trees. The female also sports a gay patch of sexual skin that puffs
8 S. L. Washburn: “Ischial Callosities as Sleeping Adaptations,” AJPA, Vol.
15, No. 2 ( 1957). PP- 269-76.
14° The Order of Primates
out and becomes excessively vascular, as a signal of invitation, be-
fore ovulation.
Listed in the same genus are the drills ( Papio leucophaeus)
— large, nearly tailless animals that roam the forest floor in West
Africa, and the mandrills ( Papio sphinx ) of Gaboon, which sally
out of the forest from time to time to raid the savannah country.
The drill has a dark green coat, a black face, and a pink posterior.
The mandrill is dark brown, with a bright red nose and a blue,
longitudinally ridged swollen area over each cheek, and a red
and blue region to the rear, a combination callosity and sexual
skin.
In a genus of its own, Theropithecus, is the gelada, an excep-
tionally large baboonlike primate, possibly of independent origin,
inhabiting southern Ethiopia. A huge mane covers its forequart-
ers, but its chest and underside are bare. These parts are as pink
as its posterior. This is the only primate beside man, apparently,
that has both a mane top-forward and bare skin elsewhere.9
These terrestrial animals are all omnivorous, intelligent, aggres-
sive, well organized in packs for concerted action, and dangerous
to man if provoked, and to carnivores. Washburn saw a pack of
baboons which were feeding in and about a herd of impala drive
off three cheetahs by simply looking threatening while the impala,
apparently confident in the protective power of their companions,
whom they served in turn as lookouts, grazed unconcerned.1 The
inference is that life on the ground, particularly in open country,
has created a situation favorable to increased versatility, adapta-
bility, and general intelligence in primates, as shown by the per-
formance of the series from macaque to gelada.
The Anthropoid Apes
Less versatile, less adaptable, more specialized, and
probably even more intelligent are the anthropoid apes, including
two genera of small ones, the gibbons and siamangs, and three
genera of large ones, the orangs, chimpanzees, and gorillas. Most
authors include them all in a single family, the Pongidae, rating
8 A comparable coat combination is seen in a terrestrial bird, the ostrich.
1 Washburn: op. cit., pp. 273-4.
The Anthropoid Apes 141
the first two as a subfamily, the Hylobatinae, and the last three as
another, the Ponginae. Both Fiedler and Straus, however, catego-
rize them as two families, Hylobatidae and Pongidae, because of
the separation of the gibbons and siamangs from the other three
both anatomically and in probable line of descent.
In either case, all apes have certain features in common. They
all have large, interlocking canines on both jaws, so that they can
chew only up and down. In the upper jaw there is a gap, or
diastema, between each canine and its neighboring lateral incisor.
This gap makes room for the lower canine when the jaws are
closed. In the lower jaw the first premolar is long, narrow, and
ridged. The outer edge of this tooth is sharpened against the up-
per canine, with which it acts as a pair of scissors.
The ape molars are not bilophodont, but retain separate cusps.
The upper and lower molars meet in different fashion from those
of the Cercopithecidae; in the monkeys the upper and lower teeth
mesh alternately, with the anterior loph of each lower tooth fill-
ing the gap between its opposite number and the one in front of it,
whereas in the apes the uppers and lowers meet more nearly
squarely on. Furthermore, the cusps of the lower molars, of all
genera of apes have the Dryopithecus pattern, named for a fossil
ape to be described presently. This is also called the Y-5 pattern,
because a tooth of this kind has five cusps, two forward and three
aft, and the grooves separating the cusps assume a pattern resem-
bling the letter Y, with its tail to the front. Much has been made
of this feature by paleontologists, who have little else but teeth to
go on.
Although one of the apes, the goi'illa, spends as much time on
the ground as baboons do, all the anthropoid apes are anatomi-
cally adapted for arboreal life through brachiation, the mode of
progression independently acquired by the spider monkeys in the
Neotropical region. In all five genera the upper limbs are longer
than the lower; both hands and feet are to some degree prehen-
sile; and the tail is gone, leaving even less of a vestige than in
man. The gibbon and siamang have ischial callosities, like the Old
World monkeys, but the three great apes sit on enlarged sphincter
ani muscles which look like portable doughnuts. All five have the
single-disc type of placenta.
142
The Order of Primates
The Gibbons: Symphalangus and Hylobates
The Hylobatidae are long-limbed and spidery, with the
longest legs in proportion to body height of any higher primate
save man, and the longest arms of all, relatively speaking, with es-
pecially long forearms. They are by far the best brachiators
among the anthropoids, but they vary their gait by running erect
along horizontal branches with their arms extended, like high-
wire artists with balancing poles. On the ground they run in ex-
actly the same way. When crossing streams too wide for aerial
jumping they swim, using the breast stroke. Compared to those of
man and the other apes, their hands and feet are long and nar-
row, and their thumbs are short and set far back in the palm, so
that the thumb tips fail to reach the first row of knuckles. This,
like the thumblessness of the spider monkey, is an adaptation for
fancy brachiation. Their teeth are small, but their canines are
long and can inflict a nasty wound. Generally, however, they
avoid direct territorial fights and instead have shouting matches,
emitting liquid calls that can be heard over a mile from an in-
flated throat pouch. All of them have small, hard, rather than
puffy, ischial callosities. There is no visible sexual dimorphism in
any of the species, and as the genitals are small and shielded by
hair, the only way to distinguish male from female in the field is
by noticing either the nipples of a female that has borne young, or
who mounts whom.
Each genus is of a different size. Symphalangus, the siamang,
has an arm span of six feet and weighs about 24 pounds. Hylo-
bates, the gibbon proper, weighs between 11 and 15 pounds, ac-
cording to the species. Both genera fall well within the size range
of the Old World monkeys, which all three pongid apes, as well as
man, exceed. Like man, the siamang has a web between the first
joints of the second and third toes and for this trivial reason he,
and not man, has drawn the name Symphalangus, presumably to
distinguish him from the gibbon. His abundant coat is a drab
black to brown, and his home is the island of Sumatra, which he
shares with two more colorful apes, the orang and one species of
gibbon.
M3
The Orangutan
The latter, which occupies a wide range in the Oriental region,
is divided into at least three species, a valid division because in
places pairs of them are sympatric. Westernmost and northern-
most is the hoolock, a small animal which is usually black but
varies through a gamut of coat colors from browns to gray and
cream. All, however, have a white bar across the eyebrows — their
distinguishing feature. Races of the hoolock ( H . hoolockii ) range
over Bhutan and Assam and up the gorges into Tibet; they cross
Upper Burma and northern Siam into Indochina and the island of
Hainan.
South of the hoolock, from Burma eastward, lives the white-
handed, or lar, gibbon, which ranges to the tip of the Malay pen-
insula. The color of its coat is also variable, but all of them have a
white ring around the face and white hair on the hands. The
skin of the face, which is bare, and of the palms and soles, is black.
To the East, in Indochina, lives the black gibbon or concolor. A
fourth group comprising one or more species is called the agile
gibbon in Malaya and the silvery gibbon in Java, and a related
form lives alongside the siamang in Sumatra. They have no special
mark, but some of the latter have bodies of one color and limbs of
another. This genetic patterning in hair color, although common
enough in mammals in general, including the lower primates, is
absent in the three great apes and in man. He, though individu-
ally and racially variable, has a single pelage color, save for an oc-
casional blond or red mustache with brown head hair, and the
graying of age.
The Orangutan (Pongo)
The orang lives in Sumatra and Borneo. Despite a separation
since the flooding of the Sunda Shelf in the late Pleistocene, the
two island races are still one species, P. pygmaeus. He is a large
animal, the adult male averaging 165 pounds and reaching 200
pounds; as the females average only 81 pounds, there is a sexual
dimorphism of 2 to 1 in weight. Sexual dimorphism is also evident
in the excessive growth of the jaws and teeth, particularly the
canines, in the adult male. The thrust of the bite is carried up-
ward through the lateral facial bones to the forehead bones to
144
The Order of Primates
the forehead in a single line without the need of brow ridges as
braces, but so strongly are the temporal muscles developed that
they meet on the top of the skull, which grows a sagittal crest to
part them and give them room for attachment. A separate trans-
verse crest at the rear supports the ends of the neck muscles
needed to counterbalance the animal’s deep, heavy jaw.
The orang’s arms are the longest in relation to his body length
of the three great apes, and he can move his legs to the side at
a right angle to his trunk, because he lacks a tendon on the hip
joint needed for erect posture. When he walks on the ground, he
uses his arms as crutches, swinging his legs between them. Like
the gibbon, he has a laryngeal sac, but instead of using it for
music-making he converts it into a pneumatic shock absorber in
brachiating. This specialization he shares with the other two great
apes. The sac extends, in the adult animal, to the shoulders,
either of which has to bear the total weight of the animal, as
much as 200 pounds in his case and more in that of the gorillas,
when he hangs by one arm. In man this laryngeal sac sometimes
turns up in heavy vocalizers, for example, opera stars and
muezzins.2
On his hairless cheeks the adult male orang sports a pair of
coarse, fleshy flanges which look like the sidepieces of a baseball
catcher’s mask and which may serve as armor against biting, al-
though this has not been proved. The orang’s hair, which is red-
dish and very long, protects his light-brown skin from the torren-
tial tropical rain. At night the orang, like the chimpanzee and the
gorilla, sleeps in a nest of his own fabrication, and in it he likes to
cover his head with leaves. He neither needs nor has ischial cal-
losities.
The Chimpanzee (Pan)
This, the most familiar of primates, frequently seen on tele-
vision and the stage, inhabits various parts of the tropical forests
of Africa from Gambia to Uganda and south to Angola. The male
2 Jane Enzmann: “The Structure and Function of the Laryngeal Sacs of the
Chimpanzee, Gorilla, and Orang-Utan,” AJPA, Vol. 14, No. 2 (1956), pp. 383-4.
i45
The Chimpanzee
is smaller than the male orang, with a mean weight of no pounds
against the orang’s 165 pounds, but the female chimpanzee is
slightly and probably not significantly larger than the female
orang, with a mean weight of 88 pounds against 81. The weight
ratio between the sexes is therefore only 1.3 to 1, which still gives
the chimpanzee greater sexual dimorphism in weight than is
found in most races of man. The stature of this animal ranges
from five feet to five feet seven inches ( 152-170 cm. ) for the
males and four feet to four feet three inches for the females
( 122-130 cm. ) . This is well within the range of human races in
the male, and well below it in the female. Morphologically
speaking, the sex dimorphism of the chimpanzee is not great, and
little difference can be found in the skull and teeth. The male
does not ordinarily have a sagittal crest.
Probably because more chimpanzees have been observed than
orangs or gorillas, we know that they are extraordinarily variable
in skin color, running from a grayish pink that is almost white to
black, with several yellowish shades between. Their color range
is essentially the same as in the races of man, and they all live
within one environmental realm. The hair color, in the blacks and
browns, is less variable, but the amount of hair on the body and
head varies greatly. Baldness is sometimes present, and the ani-
mals gray with age. The presence or absence of ischial callosities
is also variable; callosities are present in 36 per cent of the ani-
mals noted. As the chimpanzee, like the orang and gorilla, builds a
nest, he has no more need of these pads than they do.
Specialists who have tested chimpanzees — and who have even
reared some of them in their homes alongside human babies — find
that they vary enormously in intelligence and in temperament al-
though, compared to orangs and gorillas as a species, they may
be characterized as gregarious, noisy, inquisitive, and provoca-
tive. Up to the age at which the human infant learns to talk, they
are apparently our equals.
Taxonomically they are commonly divided into two species,
Pan satyrus and Pan schweinfwthii, but Schultz and others who
have studied them carefully find the division unjustified. Indi-
vidual variations within populations are at least as great as re-
146 The Order of Primates
gional ones. Moreover, there is no evidence to support the idea
that they will not interbreed, in or out of captivity.
Like the galago, the marmoset, and man, the chimpanzee has a
pygmy form. It was first described in 1929 by Schwartz, who
classified it as a subspecies, Pan satyrus paniscns. Coolidge, de-
scribing an adult specimen in 1933, called it Pan paniscns, giv-
ing it full species rank; 3 and in 1954 Tratz and Heck called it a
genus, Bonobo* In the opinion of other primate anatomists the
subspecific designation may be the correct one, because the
musculature of the two animals is the same,5 and because the two
forms are allopatric. The pygmy chimpanzee lives only south of
the Congo River, where the full-sized animal is not found.
The pygmy is about half the size and weight of the full-sized
chimpanzee, and it is morphologically pedomorphic, in contrast
to the other races and species of great apes, which are essentially
gerontomorphic. Whereas the full-sized chimpanzee has large
cupped ears, its pygmy relative has small ones like those of a go-
rilla, and a gorillalike, relatively prominent, external nose. The
pygmy has a different call from that of the full sized chimp; he
calls Hi! Hi! instead of Ho! Ho! In 1923, six years before the
existence of this race was known, Robert Yerkes acquired an
animal which had been captured in pygmy-chimpanzee territory
and which undoubtedly was a pygmy. After a little over a year, it
died, probably early in its fourth year of life. In Yerkes’s stated
opinion, it was a simian genius and the brightest ape he had ever
known, with an excess of boldness and curiosity.6 How it would
have developed as it grew older we can never know. There is one
currently in the Philadelphia zoo. The taxonomic disposition of
this animal is of special interest to anthropologists, because man
too has pygmy races, as different from full-sized people as a
pygmy chimpanzee is from his neighbors across the river. We do
not consider our Pygmies a genus, however, nor even a species.
3 H. J. Coolidge, Jr.: “Pan paniscns. Pigmy Chimpanzee from South of the
Congo River,’’ AJPA, Vol. 18, No. 1 ( 1933), PP- i-57-
4E. Tratz and H. Heck: “Der afrikanische Anthropoide ‘Bonobo,’ eine neue
Menschenaffengattung,” SM, Vol. 2 (1954), pp. 97-101.
0 R. A. Miller: “The Musculature of Pan paniscus,” AJA, Vol. 91, No. 2 ( 1952),
pp. 183-232.
6 R. M. Yerkes: Chimp Intelligence and Its Vocal Expressions (Baltimore:
Williams and Wilkins Co.; 1925).
The Gorilla
147
The Gorilla (Gorilla)
The gorilla is divided into two noncontiguous geographic
populations: the lowland gorilla of the Gaboon and Cameroons,
and the mountain gorilla of the eastern Congo west of Lakes Ed-
ward and Kivu, where it ranges in altitude from 7,500 feet to 12,-
000 feet. The habitat of the lowland animal is ordinary tropical
rain forest. The home territory of the mountain population is cool
and alpine in places, with an abundance of bamboo, the shoots of
which are a favorite food of the gorilla. The lowland form has tra-
ditionally been given a specific name, Gorilla gorilla, and the
mountain form another. Gorilla berengii, but despite certain
marked differences, as in foot form, the current tendency is to
consider them geographical races of a single species and call them
by the former name.
In both races the stature of adult males ranges from about five
feet to about six feet (152-183 cm.), although individuals of six
feet six ( 198 cm. ) have been found. This is the human range, ex-
cluding pygmies. Probably the mean weight of adult male gorillas
is about 400 pounds. Grossly heavier weights, reaching more
than 600 pounds in individual zoo and circus animals, are the re-
sult of obesity caused by overfeeding and lack of exercise; human
beings can reach 900 pounds under similar circumstances. The
adult female is much smaller, probably weighing not more than
200 pounds. The weight ratio between the sexes, then, is about 2
to 1, as among the orangs.
Sexual dimorphism is exaggerated in the head and face. The
skull of the male gorilla has enormous teeth, particularly canines,
and huge brow ridges, sagittal crest, and nuchal crest, although
these flying buttresses vary greatly among individuals in size and
form. The considerable size of the brain case is thus overshad-
owed by the overdevelopment of the masticatory apparatus and
its dependent braces. The female skull, however, can hardly be
distinguished from that of a male chimpanzee. Gorillas almost
never have ischial callosities. Their skin is usually black, their ears
small, and the soft parts of the external nose stand out from the
facial plane.
148
The Order of Primates
Most of the differences between the two races of gorillas are
trivial compared to those which separate human subspecies, and
concern relative trunk and limb lengths and skull form. However,
the difference in foot form does suggest that they may be two
species. Although the foot of the lowland gorilla is broad and
short by simian standards, its great toe is pointed at an angle from
the axis of the foot, whereas in the mountain gorilla the great toe,
although still short, is in line with the foot, like that of a man.
Like the chimpanzee the gorilla sleeps and sometimes feeds in
trees, but he moves from one tree to another by climbing down
and walking from trunk to trunk on the ground; and in the bam-
boo forest he habitually feeds on the ground. Like the other great
apes he walks on all fours, on the outer edges of his feet and on
the knuckles of his hands. He can stand erect when he wishes,
but when moving fast, like the others he runs on all fours.
The gorilla is less communicative than the chimpanzee and al-
most always silent; he is less aggressive in the wild, but sometimes
more so in captivity.7 What noise he makes he produces mostly by
drumming on his chest with his fists. Which of the two apes is the
more intelligent is a matter of controversy. Gorillas are the most
difficult apes to rear in captivity, and the least communicative.
The chimpanzee wears his intellectual heart on his sleeve, so to
speak; the gorilla’s is deeply concealed.
The Hominidae (Homo)
Represented, like each of the three Pongidae, by a single
living genus and species, the family of Hominidae differs from its
fellow giant catarrhines in a number of easily enumerated fea-
tures. Man’s brain is about three times as large as those of the
great apes, although at birth it is smaller than the orang’s. The
human brain is also more specialized than those of the others.
Man’s teeth are generally smaller. His canines in particular are
smaller. They are no longer than the incisors and premolars that
7J. T. Emlen, Jr.: “Current Field Studies of Gorillas,” CA, Vol. 1, No. 4
(i960), p. 332.
Emlen: “In the Home of the Mountain Gorilla,” AK, Vol. 63, No. 3 (i960),
pp. 98-108.
The Hominidae
x49
flank them, so that they do not overlap. Consequently there is no
diastema, or gap in the gum line, between the canines and the
lateral incisors on the upper jaw nor one between the canines and
the first premolars on the lower.
Whereas all the apes are specialized for brachiating, man is spe-
cialized for walking erect, with consequent differences in the
spinal column, pelvis, upper and lower limbs, hands, skull, jaw,
and vocal apparatus. As he neither sleeps in trees nor has yet be-
come anatomically adapted to a life in chairs, man has either lost
or failed to develop ischial callosities. Most men are less hairy than
most apes, and in some races man is virtually hairless, except on
the head, where he sports a mane, and in the axillae and pubis. In
some races the male also has a beard. His skin color covers the pri-
mate range, even to the blue of the drill’s nose, which is matched
by the so-called Mongolian spot on the skin over the sacrum in
some races, and the blue penis of some South American Indians.
His hair color is less variable because it lacks greens, and his
eye color is possibly more variable, with an emphasis on blues,
grays, and greens as well as various browns and intermediate pat-
terns. His hair form is more variable than that of all other pri-
mates; only man has the woolly or spiral hair of Negroes and
Bushmen.
In his tolerance of extremes of climate, in the variety of terrain
he is able to inhabit, in his general adaptability, aggressiveness,
and ability to live in groups, man more closely resembles the mon-
keys who live on the ground, particularly the macaques, than he
does any forest-bound ape. The human populations which live
most simply differ little, if at all, from other primate populations in
their relationship to other species of animals and to the landscape.
Although partly carnivorous, they kill only what they need to eat,
and leave nature in balance as they find it. Now and then they
burn over hunting grounds, and this action favors some plant
species over others and provides certain browsing and grazing
animals with more food than before, but self-ignited forest fires
have the same results. Still, it is possible that in various non-
tropical regions, particularly in the Palearctic and Nearctic, hu-
man hunters may have caused, or at least hastened, the extinc-
tion of several species of mammals, mostly oversized ones such as
1 50 The Order of Primates
the mammoth in Europe and Asia, and the mastodon, giant sloth,
and even the horse, in North America. Ecological disturbances
caused by man may have been more substantial at the hunting
level in cold climates than in warm ones, particularly the wet
forests.
Nevertheless — setting aside these extinctions for the moment —
it was only about 8,000 years ago that man began to enlarge his
ecological niche by doing what certain invertebrates, the ants and
termites, had been doing for millions of years — building elaborate
houses, domesticating other species and raising vegetables, stor-
ing food in great quantities, and grouping themselves in elabo-
rate hierarchies. As all human populations do not share modern
civilization, some peoples, fast disappearing, are still in the same
ecological balance as lower primates, whereas others, rapidly in-
ceasing, have already hit the moon and are orbiting men around
the earth.
ss
5
MAN’S PLACE AMONG
THE PRIMATES
r/ie Bearing of Primate Studies on Racial Origins
-L -J ver since Darwin, there has been a diversity of
opinion about the position of our species among the primates.
Darwin himself considered us closest to the great apes, and Hux-
ley agreed with him. Mivart, however, argued that the Old World
monkeys are our closest kin. Later on, Wood-Jones bypassed all
apes and monkeys impartially and maintained that man had de-
rived from tarsiers. Today Gregory, Schultz, and Washburn favor
the apes, and Straus supports certain aspects of Mivart’s earlier
position.
Lacking special training or experience in primatology, I am in
no position to argue on one side or the other. In fact, a question
arises as to why, in a book on the origin of human races, this sub-
ject has to be discussed at all. The answer is simply that we need
all the evidence we can get to solve the problem of our descent,
which is inadequately documented paleontologically. If we know
to which other living primates we are most closely related in
anatomy, physiology, and behavior, we shall be in a better posi-
tion to interpret the evidence of fossil bones than if the bones were
all we had. This will help us determine the time at which our an-
cestors and theirs parted company. Such information will, in turn,
give us some idea as to the time when the geographical races of
man could have begun to differentiate. Finally, comparisons with
other primates may help us decide which specializations of living
human races are old and which are new.
152
Man’s Place among the Primates
To Brachiate or Not to Brachiate
The most conspicuous difference between man and the other
primates is our erect posture, with bipedal locomotion and free
hands. Although not the only criterion of genetic relationship, it
has become the most controversial. Most orders of mammals have
one principal method of locomotion each, be it swimming, flying,
running about on the ground, or climbing trees. But the single or-
der of primates has a wide range, including climbing, hopping,
tail-swinging, walking on all fours (quadrupedal), brachiating,
and walking on two feet (bipedal). In an early primate stage,
man’s ancestors must once have been arboreal and quadrupedal.
Whether or not they brachiated for a while before becoming bipe-
dal is the moot question. Anatomists have argued it back and
forth, bone by bone and muscle by muscle, largely motivated by a
desire to find out, in terms of this single feature, if man is or is not
descended from some kind of ape.
This concept is based on a false premise — that if man’s ances-
tors once brachiated, he must be descended from early, brachiat-
ing apes, and conversely, that if man’s ancestors did not brachi-
ate, he is not descended from early apes. It is false because
brachiation can be acquired in different primate lines independ-
ently. Spider monkeys, langurs, and apes, belonging to three dif-
ferent superfamilies, have all adopted this specialized means of
locomotion. If man’s ancestors did indeed brachiate at one time,
they may have done so independently of the ancestors of the apes.
This we shall explore in the following chapter when we examine
the fossil record.
To my mind the essence of the argument, at this point, is not to
trace the descent of man nor the history of his locomotion, but to
explain the freeing of the human shoulder girdle, arms, and hands
from whatever kind of locomotion man’s remote ancestors prac-
ticed. The human upper extremity is unique among bipedal ver-
tebrates in that it is neither involved in locomotion, except for
guiding animals and steering vehicles, nor degenerate. Ostriches,
emus, rheas, and other flightless birds have tiny degenerate wings
covered with pin feathers or down. Kangaroos, jerboas, kangaroo
rats, and most other nonarboreal, bipedal, jumping mammals have
The Bearing of Hominid Teeth on the Erect Posture 1 53
tiny, mostly useless, arms and hands. So did the bipedal dinosaurs.
Man s arms are at least as long, in ratio to body length, as those of
quadrupedal monkeys, and man’s hands at least as large and as
mobile as theirs. Why?
Two theoretically reasonable explanations come to mind, each
involving a direct shift from one function of the forelimbs to an-
other without time off between for degeneration.
( 1 ) Descent from Brachiators. Our remote arboreal ancestors
began to brachiate and specialized to a certain extent, say, as far
as the woolly monkey has done. This gave them a powerful upper
extremity. Their hands did not become specialized for hanging,
like those of the great apes, because our ancestors were then small
animals too light to need this adaptation. When a brachiating ani-
mal is moving through the trees, and also when it is seated, the
axis of its trunk is vertical to the ground. Such au animal, when it
comes to live on the ground, can retain this vertical position only
by walking erect. As the weight of the entire body falls on the
pelvis and legs, instead of on the arms and shoulder girdle, the
pelvis and legs had to become adapted for bipedal walking, par-
ticularly as the body grew heavier, whereas the arms and hands
were ready-made for the use of tools and weapons, carrying in-
fants, and other manlike and womanlike activities.
(2) Descent from Quadrupeds. Let us suppose that instead of
brachiating our remote ancestors left the trees while they were
still quadrupeds, like the prosimians and most of the monkeys.
When they got to the ground they walked about on all fours, like
the macaques and baboons. Before they could acquire long snouts
like baboons or bright buttocks like drills, they began, for some
reason or other, to stand up and walk like men. As there are inter-
mediate stages of standing and walking erect, our ancestors could
have done this gradually, just as other primates are becoming
adapted to brachiation gradually, and just as the ancestors of bats
must have glided like flying squirrels before they learned to fly.
The Bearing of Hominid Teeth on the Erect Posture
Whether they had been brachiating or not, what caused
our ancestors to find the erect posture advantageous? The answer
may be related to the peculiar form of their teeth.
154
Man’s Place among the Primates
All primates use their hands for picking food off the stem, off
the ground, out of the nest, or wherever, and for peeling, break-
ing, or otherwise processing it, and for putting it in their mouths.
Unlike many other animals, they neither graze nor browse. Flight-
less birds peck their food with their beaks, and kangaroos bite off
theirs with their teeth. Most of the primates, however, have long
canines, with which they do a bit of food processing. Holding a
tough-skinned fruit in one hand, the monkey or ape will rip off its
husk with his canine teeth. With these same canines they fight, if
they cannot win by scowling and bluffing. It is in order to succeed
at fighting, not to process fruit, that males have longer canines
than females.
Like some of the smaller monkeys, the hominids have short
canines set in line with their other teeth. Because of this handicap
they have to use both hands at once to peel a tough fruit. As this
leaves no hands at all for support in the trees, it brings the animal
down to the ground. Because of it they are also obliged to use
their hands in fighting — and it is difficult to fight with your hands
when on all fours. The dental eccentricity of hominids thus favors
both life on the ground and life standing up. Our peculiar teeth
may not explain all the whys and wherefores of the erect posture
and powerful forelimbs of man, but they can easily have influ-
enced this combination.
A Few Details of the Posture Story
The anatomical evidence for and against the belief that
our ancestors once swung hand over hand in the trees is long and
involved and serves the purposes of this book only in documenting
the fact that some races seem more arboreally constituted than
others. This will be a very short summary of the primate evidence,
and man will here be treated temporarily as a homogenous spe-
cies.
Like most other four-footed animals, the nonbrachiating, sub-
human primates hold their vertebral columns horizontally, sling-
ing their internal organs downward from them by the force of
gravity. Owing to the usual mechanical predominance of the hind
over the forequarters, the vertebral column has a particularly long
A Few Details of the Posture Story 155
lumbar and a particularly short thoracic or rib-bearing segment.
One vertebra, known as the anticlinal, which is generally the
tenth thoracic, acts as the center of a spring. It is the continental
divide, so to speak, between the muscles of the hind and fore-
quarters; and the dorsal spines of the vertebrae in front of it point
backward and those behind it point forward. This arrangement is
ideal for jumping, with the bulk of the thrust coming from the
rear, as is the case with most other four-footed mammals.
All four limbs are usually flexed; in many species they cannot
Fig. 5 The Jumping Skeleton. A climbing, jumping, quadrupedal primate: the
marmoset Hapale. Note the dorsal spines of the vertebrae. The ninth thoracic
vertebra acts as fulcrum, and its spine points upward. Those in front point
backward, those behind point forward. ( Drawing after Gregory, 1951. )
be held out perfectly straight at the elbow and knee. But seen
from the front and rear each extremity seems to form a straight
column, as it does in dogs, cows, and most other quadrupeds, and
the two hands and two feet are a considerable distance apart
when in motion. The rib cage is oval in horizontal section, with
the long axis running from back to front; the thoracic vertebrae
are set at or above the upper rib level; the sternum is narrow; and
the scapulae are placed alongside the rib cage. The legs are longer
and stronger than the arms, and the thumbs are well developed.
156
Man’s Place among the Primates
Fingerprint ridges of the kind made famous by the F.B.I. com-
pletely cover the skin of the palms and soles in the Old World
monkeys, whereas in the New World monkeys and prosimians they
cover only parts of the surfaces. New World monkeys that hang
by the tail have similar ridge patterns on the business surface of
that organ as well. In the Old World monkeys the ridge patterns
of the palms and soles are simple. Although their hands have con-
siderable mobility, as do the hands of all primates, in the wrist the
carpal bones articulate with both radius and ulna, and this pre-
vents them from rocking the hand from side to side as hitchhikers
do when thumbing rides. When traveling on all fours, the Old
World monkeys place their palms down, with fingers extended, on
the ground.
As all monkeys, and even some of the prosimians, sit up when
resting or sleeping, the erect posture is habitual when they are
not in motion. Consequently the neck is moderately long and
flexible and the skull is hafted to the cervical vertebrae in a com-
promise position, so that the animal can look ahead when the
trunk is either horizontal or vertical.
In the bodies of the brachiating apes an entirely different type
of adaptation is found. Compared to the quadrupedal monkeys,
their vertebral column is longer in the rib section and shorter in
the lumbar region. It no longer serves as a spring for jumping,
but simply as a vertical rod for holding the body’s weight. It is
somewhat like a pendulum. As most of the locomotion is per-
formed by the arms, the anticlinal position has moved down about
three places, and now is in the first lumbar. The muscles of the
shoulder girdle have also advanced downward, and those of the
lower extremities have retreated in the same direction. Some of
the shoulder muscles now reach down to the rim of the pelvis.
In the monkeys, as in many more primitive vertebrates, the
muscles tend to be attached directly to the long bones, which
are ridged, lumped, and pitted to receive them. But in the brachia-
tors the same muscles join the bones indirectly through the inter-
mediacy of tendonous sheaths, called aponeuroses, which leave
the surfaces of the bones tubelike and smooth. This arrangement
greatly increases the mobility of the limbs. Also, the limb muscles
have relatively short, fleshy “bellies,” as the main bodies of the
A Few Details of the Posture Story 157
muscle bundles are called, and relatively long, tough tendons.
This combination is best suited to withstand the shock of the
whole weight of the falling body as it is caught up short by the
muscles of a single arm in brachiating. The shock is further
cushioned in heavy adult apes by the laryngeal air sac.
In section the brachiator’s rib cage is shaped like a heart with-
out a point at the rear, and flat in front; the ribs arch behind the
vertebrae, and the whole enclosure is relatively broad and flat, as
is the sternum. The scapulae (shoulder blades) are set behind the
rib cage, not beside it. The arms are long and mobile at shoulder
and elbow, and the arm can be straightened out completely at the
elbow or even bent back, as in the gorilla.
In our own bodies the legs do not form straight columns, dog-
Fig. 6 Transverse Rib-cage Sections of Jumpers,
Bhachiators, and Man. ( Drawings 1 and 3 after Hamil-
ton, 1956; drawing 2 after Raven, 1950.)
and monkey-wise, when seen from the front or from behind. Our
thighs converge from hip to knee, and our lower legs run parallel
to each other from knee to ankle. The angle between the axes of
the upper and lower leg portions is called the carnjing angle.
It keeps our trunks from swaying from side to side as we walk. It
is an adaptation to the erect posture.
According to the same principle, the apes need a carrying angle
at the elbow. Their upper arms converge from the shoulder and
their lower arms run parallel to the wrist. When an ape is brachiat-
ing, each hand in turn passes directly over his head, and his body
is thus kept from swinging back and forth sidewise.
Although they do not walk bipedally, the apes also have carry-
ing angles in their legs. This allows them to bring their feet to-
1 58 Man’s Place among the Primates
gcther in parallel fashion when grasping a straight object, such
as the limb of a tree. Although we do not brachiate, we have
carrying angles in our arms. This allows our arms to clear our
hips, which is particularly useful to women, and it enables us
to grasp a straight object with both hands, just as apes do with
their feet.
Fig. 7 The Carrying Angle in Apes and Men. The carrying angle is the
angle formed at elbow and knee in brachiators and man. In an animal that moves
itself with one pair of limbs at a time — anns or legs — this angle is needed to keep the
body upright. It is not found in four-footed animals which walk or run with at
least one limb of each side on the ground at all times, or when the body is
entirely in the air, as in leaping or hopping.
The brachiator’s arms are longer than his legs, and his forearm
is especially long. His wrist bones articulate with the radius only,
and his hand can be rocked sidewise, which is very useful in
brachiating. In the three large apes the fingers cannot be straight-
ened out when the wrist is bent forward; this is due to short,
strong tendons in the fingers that make the hands into hooks and
A Few Details of the Posture Story 159
permit the animal to hang by the hands for long periods without
muscle strain.
As a price of brachiation the thumbs of the apes have become
short and imperfectly opposable. The great toe, in compensation,
is partially opposable except in the mountain gorilla, whose feet
are to a certain extent adapted to walking on the ground. In all
the apes the ridge patterns of the palms and soles are relatively
complicated. When walking on the ground, the three large apes
walk on the backs of their fingers; the gibbons do not ordinarily
touch their hands to the ground.
The ape skull is hafted more or less as in the Old World mon-
keys, but owing to the exaggerated weight of the teeth and jaws
the neck muscles creep higher on the neck than in most monkeys.
The neck itself is short and not very mobile.
The lumbar region of the vertebral column is longer and heavier
in man, the bipedal primate, than in the apes. Unlike the compar-
able structures of either apes or monkeys, his vertebral column
has an S curve when seen from the side, and the center of gravity
of his body passes through this bony column vertically. His pelvis
is relatively short and broad, and his ilium (hip blade) is particu-
larly short. As among the apes, the muscles of his shoulder girdle
reach far down the trunk — in some cases to the pelvis — and the
muscle attachments tend to be made directly on the bone, as
among monkeys.
In the ischium, the most distal and dorsal (hindmost and rear-
most ) of the three bones that fuse to form the os coxae, or pelvic
bone, man differs from both monkeys and apes. Because when
man walks he jolts his visceral organs downward with each step,
his lower pelvic bones must converge inward as far as possible to
support them, while still allowing space, in the female, for the
birth canal. Between the bones stretches a firm web of ligaments
and muscles. If this network fails, the result is a hernia. In the Old
World monkeys the pelvic opening points backward, and in the
apes the shoulder girdle takes the strain of locomotion and cush-
ions the blows to the viscera.1
These differences in locomotor habit have left the Old World
1 H. O. Elftman: The Evolution of the Pelvic Floor of Primates,” AJAn ,
Vol. 51, No. 2 (1932), pp. 307-46.
160 Man’s Place among the Primates
monkeys and apes with an ischium that is long and flares out-
ward, whereas in man the ischium is short and bent inward. The
monkeys and gibbons sit on callosites covering the ends of these
bones; in the great apes the lower part of the ischium serves as an
anchor for the gluteus maximus muscle, which does not cover it
but extends to the side.2 In man the gluteus maximus covers the
ischium, helping to hold the trunk erect and forming a buttock,
which is unique among primates.
Monkeys do not ordinarily move about bipedally. When apes
do they run to keep from falling forward; they can stand erect
without moving only a very short time, as when a male gorilla is
Fig. 8 Feet of Apes and Men. ( Drawings after Schultz, 1956. )
drumming on his chest. Standing erect continuously and true
walking are exclusively human attributes.
Although man’s arms are shorter, relatively, than those of the
apes, they are fully as long, compared to body length, as those of
quadrupedal monkeys. They are also mobile. Like apes’ arms,
they too hang at a carrying angle, and their wrist bones also
articulate with the radius alone, allowing the hand to be rocked.
However, the thumb is as long as or longer than the monkey’s
thumb. Also, the hand can be extended completely, and the thumb
can be opposed to all four fingers. A baby crawling or walking on
all fours sets the palm of the hand down flat, with fingers ex-
tended, just as a monkey does. The palm- and sole-ridge prints
2H. C. Raven: The Anatomy of the Gorilla (New York: Columbia University
Press; 1950), p. 57 and figures.
A Few Details of the Posture Story 161
are simple, like a monkey’s rather than an ape’s. The legs are
uniquely long. Man has relatively immobile knee and ankle joints,
with large, flat-faceted tarsal bones, and a long great toe, in line
(when not deformed by shoes) with the other four.
The head of the bipedal primate is hafted more anteriorly than
in any other primate. His neck is longer than that of apes, and the
animal can look ahead without peering over the tops of his glasses,
so to speak, only when he is standing or sitting in an erect posi-
tion. Probably because his assumption of the erect posture has
moved the female genital organ considerably forward, only he
and the pygmy chimpanzee among primates copulate ventrally,
like the porcupine, who does it for a different and even better
reason.3
In evaluating these perplexing and somewhat contradictory
comparisons we must remember that four closely related families
of animals have become adapted to four principal means of loco-
motion: quadrupedal in trees, quadrupedal on the ground, bra-
chiating, and erect bipedal. This is a wide range for so small a
group. In order to achieve this degree of differentiation, the
genetic capacities of these families must have been exploited to
the utmost by differential growth rates. Neoteny, pedomorphism,
and gerontomorphism reach their peak in man and in his close
primate kin.
Large brachiating and bipedal primates have several functional
requirements in common, one of which is the absence of an exter-
nal tail. To both, this appendage would be useless, if not an actual
hindrance. A large ape is too heavy to hang from one, and a man
needs what is left of his to help close his pelvic opening. Its ab-
sence in the gibbons suggests that the monkeys which live on the
ground have short tails, some mere stubs, because they need no
balancing appendage. Its absence among all apes, great and
small, suggests that their common ancestors spent a prebrachiat-
ing period on the ground, rather than going directly from one
kind of arboreal locomotion to the other, as can be postulated in
the case of the New World monkeys.
Another similarity between apes and men is the shape of the
8 Other mammals said to do this are the hamster and the two-toed sloth.
Bourliere: The National History of Mammals, p. 159.
162
Man’s Place among the Primates
chest. For swinging apes and walking men, a broad, flat rib cage
keeps the center of gravity below or above, as the case may be,
the focal point of locomotion in the anteroposterior plane, and its
greater width is compensated for by the carrying angle of the
propelling limbs. Four-legged animals do not have these gravity
problems because, unless cantering, galloping, single-footing, or
jumping, they have one foot of each side and pair on the ground
at all times. Also, in apes and men alike, a broad chest extends
the animal’s reach and gets in the way least when the arms are
busy in brachiation or work. Both apes and men need, and have,
dorsally situated scapulas. As to the carrying angle, this is useful
in all four limbs to both apes and men, but in opposite ways.
This brief survey does not conclusively indicate whether or not
our ancestors brachiated. Even if it did, it would not settle the
question of how close we are in descent to either the Old World
monkeys or the apes. Other anatomical comparisons favor the
apes. All the Old World monkeys have ischial callosites. The gib-
bons have small ones, and these are also found in roughly a
third of the chimpanzees. The orang, the gorilla, and men lack
them. The Old World monkeys have two-disc placentas. Men
and the apes have single-disc ones. In these features the apes
seem to be man’s closest primate relatives.
The Evidence of Teeth
Much can be learned from the standby of the paleontolo-
gists, teeth. Human beings have the same dental formula as
both the Old World monkeys and the apes, 2 1 ,2,3, wpjcp sets
2:1:213
them all apart from the prosimians and platyrrhines. Man also
differs from the Cercopithecines on three counts and from the
apes in two of these. Both Old World monkeys and apes have a
long upper canine, separated from the first upper premolar by a
gap. In both, the first lower premolar has a shearing edge. The
Old World monkeys alone have bilophodont molars. As man has
none of these three dental distinctions, on this score the two
groups of Old World primates are closer to each other than either
is to man.
The essential feature of hominid dentition is that it enables the
The Evidence of Teeth 163
hominid to grind the lower against the upper teeth, from side to
side and from front to back, instead of just compressing the jaws
together ape-wise, up and down. Our method is highly advantage-
ous for an omnivore with carnivorous tendencies as it gives an
animal a high masticating efficiency per unit of tooth surface area
in reducing coarse food, such as skin, lean raw meat, and tendons,
as well as tough roots, into digestible fodder. Thus the reduction
of the canines, which seemed at one point a rather infantile and
disadvantageous mutation or retention, actually gave the homi-
A B c
Fig. 9 Occulusion of Canines in Apes and Hominids. In the apes (A) the
upper canine overlaps the lower first premolar and lower canine, and in biting
and chewing it touches these teeth, particularly the lower first premolar, against
which it grinds a shearing edge. In the hominids (Australopithecus and man) (B)
the unworn canine overlaps the same two lower teeth at first, but after they have
been worn down to a certain extent by a rotary motion, the points of all three are
worn oft and all of them have acquired smooth occlusal surfaces (C). As modern
civilized man does not chew enough to arrive at stage C, his teeth may remain at
stage B until death. 1
nids a distinct advantage over other primates who live on the
ground and seek the same kind of food, because it permits the
teeth to concentrate on one task only, mastication, instead of
tlnee, fighting, mastication, and peeling coarse-skinned fruit. The
change that made this transformation possible was the release of
the hands from the duty of locomotion for the work of fighting olf
rivals, killing game, peeling and even cutting up food. The teeth
of piimitive men and of adult fossil hominids are worn flat
from chewing, whereas those of apes retain their original cusp pat-
terns for life.4 Although our teeth are apelike rather than monkey-
4 Anyone civilized enough to read this book will probably find that he cannot
grind his teeth together from side to side because his upper incisors and canines
overlap his lower ones. This is not the fault of his genes, but the fault of his
parents, who fed him on soft food. He did not have the opportunity to develop
his hominid bite. *
164 Man’s Place among the Primates
like, they are really sni generis and distinctive, like the erect
posture — a hominid hallmark — and we have probably had them
for a long time.
In another respect, however, our teeth do resemble those of the
apes and Old World monkeys, and incidentally those of the
Madagascar lemurs. Seen microscopically, the enamel prisms have
straight edges and are separated only by a little insterstitial ma-
terial. In the platyrrhines, lorises, and tarsiers, the enamel prisms
have wavy edges and are separated by larger amounts of inter-
stitial material. Both forms are found in fossil prosimians. What-
ever this evidence means paleontologically, it clearly separates
the higher suborders of the New and the Old World.5
The Evidence of Embryology
S o f a r we have been considering the evidence of relationships
within the primates in terms of gross anatomy, which means the
anatomy of adult individuals. Not every individual, however, is an
adult. The comparative anatomy of individuals aged six years, for
example, is also of value, and we can learn a great deal from a
comparative study of individuals who have not yet emerged from
the womb.
These last were of particular interest to the German zoologist
Ernst Haeckel, a contemporary of Darwin, although younger.
Deeply moved by Darwin’s work, he propounded, in 1866, the
biogenetic law, also known as the law of recapitulation, the theme
of which is that ontogeny recapitulates phylogeny. This means
simply that each one of us, from fertilization to birth, passes
successively through the forms of all his ancestors, being in turn
amoeba, worm, fry, tadpole, and so on. This recapitulation is said
to take place partly in the embryo and partly in the larva, the lat-
ter being a grub in the insect world and a tadpole in the am-
phibian. Among mammals the larva is called a fetus. The embryo
becomes a fetus at the stage at which one species can be told
from another. In man this occurs at about the beginning of the
ninth week.
5 C. T. Regan: “The Classification of the Primates,” Nature, Vol. 125, No. 3143
(1930), pp. 125-6.
The Evidence of Embryology 165
Like many other laws and rules of the nineteenth century,
Haeckel’s has been variously supported and attacked. In its
liteial sense it has been generally repudiated, but the fact remains
that it is true in essence, for organisms do repeat ancestral forms,
within certain limitations.
Each organism goes through a series of the corresponding em-
bryonic forms of its ancestors, not of the adult forms. For reasons
of economy, some stages may be skipped entirely, as in certain
limbless amphibians which, although descended from four-legged
ancestors, have no trace of limbs at any embryonic stage. This
particular omission can be explained by neoteny, already dis-
cussed in Chapter 1. In addition, certain structures needed for
survival during larval or fetal life, but of no use later, appear at
that time only, without reference to general recapitulation. An
example is the twig-mimicking larval form of certain caterpillars
destined to grow into moths.
The sequence of developmental stages of each structure follows
the evolutionary order, with or without omissions, but different
structures do not necessarily keep to a single timetable. The fact
that feature X reaches a more or less adult functional form at
the end of fetal life whereas feature Y is still quite retarded, rela-
tively speaking, at time of birth does not mean that X evolved
earlier in the ancestral phylogeny than Y did. A fetus has imme-
diate postnatal survival requirements of its own and only so much
uterine space to develop in, and therefore prepares itself primarily
for urgent needs that arise at birth. Some of the structures with
lower priorities are left to complete their growth afterward, just
as soldiers about to go into battle clean and load their weapons
first, rather than shaving, or polishing their boots.
A fawn and a colt are born with long legs on which they can
run closely behind their mothers’ tails a few minutes after birth if
the mother is disturbed. A newborn gibbon, which does not yet
need to brachiate but needs immediately to cling tightly to its
mother s fur and to stay there until after it has been weaned, is
born with strong fingers and short forearms, which grow long
thereafter. By the same token, a human baby, who does not need
a big brain in the cradle but would have a hard time passing an
oversized head through its mother’s pelvic opening, is born with a
166 Man’s Place among the Primates
head even smaller than that of an orang. Like a gibbon’s forearm,
the baby’s brain grows prodigiously after birth and is ready for
action when needed.
Schultz once compared twenty-one features in three series of
fetuses of Old World monkeys, apes, and men. He considered
only diameters, which he painstakingly measured, and propor-
tions, which he calculated, at the beginning of fetal life, at what
corresponds to the sixth prenatal month in man, and at full term.
He found an extraordinary similarity between the three groups,
and some differences.6
The catarrhine primates, including Old World monkeys, apes,
and men, are gradually transformed in the fetal stage from a com-
pactly packaged embryo to a more regionally differentiated or-
ganism. However, the forearm length of monkeys and apes is
prenatally more nearly adult than in man because monkeys and
6 A. Schultz: “Fetal Growth of Man and Other Primates,” ORB, Vol. l, No. 4
(1926), pp. 465-521.
Twelve features of all three develop more or less the same way and at the same
rates, as follows. Chest circumference decreases relative to trunk height. Hip
breadth relative to trunk height increases. Hip breadth relative to shoulder breadth
increases. The position of the umbilicus moves higher on the trunk. Upper-limb
length relative to trunk height increases until the sixth month and then decreases.
Lower-leg length relative to thigh length increases. Hand breadth relative to
hand length and foot breadth relative to foot length both decrease. Thumb length
relative to hand length decreases. Average diameter of head relative to trunk
height decreases. Nose breadth relative to face breadth decreases. Interocular
breadth (distance between the inner eye comers) relative to face breadth de-
creases.
The development of four of the features takes the same direction in all three
samples, but the monkey and ape fetuses change faster than man’s do. These
changes are as follows. Forearm length relative to upper-arm length increases.
Total face height relative to head height increases. Upper-face height relative to
face breadth increases. Nose breadth relative to nose height decreases.
In two features the direction is the same in all three samples, but man changes
the faster, as follows. The upper-limb length relative to lower-limb length de-
creases. The lower-limb length relative to trank height increases until the sixth
month and then decreases.
In two features the monkeys and apes form a bloc, differing from man in the
direction of change at least in part of the fetal cycle. Whereas in man the nipples
migrate lower down on his chest until the sixth month and then move up again,
the nipples of monkeys and apes move higher constantly. Man’s head grows
narrower in proportion to its length, but the heads of monkeys and apes retain
a constant proportion or grow broader.
In only one feature does man split the simian bloc. In the apes and in man,
the chest grows broader compared to its depth. In the monkeys it does the op-
posite.
The Evidence of Embryology 167
apes need their forearms for locomotion. Also their face length
is more nearly adult prenatally because they need to chew earlier
than man does. On the other hand, man exceeds both monkeys
and apes in the early growth of the lower limbs, which must be
long for walking erect. The chests of both apes and men, unlike
those of the monkeys, undergo a prenatal broadening and flatten-
ing which anticipates both brachiation and bipedal walking,
neither of which need be derived from the other, on this evidence.
In other words, the measurement of fetuses fails to tell us whether
or not man’s ancestors brachiated. Were we, indeed, to judge
Fig. 10 Body Proportions of Newborn Primates. Note
that the orangutan is bom with the largest head. (Drawings
after Schultz, 1926. )
from this evidence alone, we could not even be sure that apes do.
Some of Schultz s morphological observations are more helpful.
The human thumb, which starts out in line with the fingers, be-
gins very early in fetal life to rotate until it comes to rest at a 90 0
angle to the axis of the fingers. At the same time its place of
origin, in the same knuckle line as the other digits, moves wrist-
ward. In the Old World monkeys and apes the process of rotation
and migration begins later and the rotation is less complete. The
human thumb remains long, whereas the thumb of the monkey
and the ape does not grow as long, particularly in the apes.
168 Man’s Place among the Primates
In the foot of monkeys and apes a comparable rotation and
migration occurs, but it does not in the human fetus. In the other
primates the longest toe is usually the third, whereas in man the
first or second eventually becomes the longest. In 4 to 5 per cent
of human fetuses at the beginning of the third month, however,
the third toe is still the longest. In both the hand and the foot,
therefore, differences in the rate of growth of the digits and in the
timing of the rotation of thumb and big toe separate the human
fetus from those of the other two categories of Old World pri-
mates. To say that one is more fetalized than the other is an over-
simplification. Rather, man’s great precultural specialization, a
combination of bipedal locomotion and manual dexterity, makes
its mark early in the fetal timetable, overshadowing the anatomi-
cal preparations for the less radical but equally spectacular feat
of his pongid cousins, brachiation. This may even imply that we
walked erect before the apes took to the trees.
Further evidence comes from the study of fetal body hair. It
has often been observed that when an animal moves into a new
medium, such as air or water, or acquires a new means of loco-
motion that will take it into new environments — and bipedal
walking is such a means — its coat changes. Thus whales, which
are known to be related to the even-toed land mammals such as
sheep and cattle,7 are hairless. Birds lost their scales when they
took to the air and developed feathers instead; flightless birds in
several parts of the world independently retained fetal down.
The arboreal primates preserve the primitive mammalian hair
coat that some of the monkeys living on the ground have partly
lost; and man, with his new means of locomotion that takes him
even farther afield than macaques, has a coat as fetal as those of
ostriches, probably because, in a wide variety of climates, par-
ticularly in hot, open sun, a nearly hairless body suits our thermal
requirements better than a hairy one does. Bolk indicates that in
the fetal life of four selected primates the following series of coat
reductions can be seen.8 Monkeys are born completely covered
with hair, as in the adult form. In the gibbon, the head and back
7P. A. Moody: An Introduction to Evolution (New York: Harper and Brothers;
1953), PP- 103-13-
8 P. R. de Beer: Embryos and Ancestors (London: Oxford University Press;
I95i). P- 58-
L. Bolk: Das Problem der Menschenwerdung (Jena: Fischer Verlag; 1926).
17° Man’s Place among the Primates
are covered with hair and the rest grows out after birth. In the
gorilla, only the head is covered and the rest grows out later. In
man also only the head is covered, although there may be a reten-
tion of fetal downy hair (lanugo), which soon disappears. Other
body hair appears later, particularly after puberty. Schultz has
remarked that some men of his acquaintance are hairier than
some gorillas, at least on the chest. This evidence suggests that
the ancestors of the apes, including the gibbons, did a stretch on
the ground before they began brachiating.
To return to Schultz’s measurements of fetuses: he found that
throughout fetal life the brain cases of monkeys and apes remain
long and narrow, whereas those of man are much more globular.
Kummer, who has studied these changes in detail, concludes that
in the form of the brain case man is not, as Bolk said and as many
others have repeated, a fetalized ape but a creature sui generis.9
Regardless of head shape, however, in the relative size of the
lobes of the cerebral hemispheres the brain of the fetal chim-
panzee resembles the adult human brain more than it does its
own adult form.1
Kummer ’s excellent drawings bring out still another embryo-
logical fact pertinent to our inquiry: the prominent nasal skeleton
of man, particularly of European man, when seen in profile, is
not a gerontomorphic feature unless excessively developed. Rather,
it is to a certain extent a fetal feature. In the human fetus the
nasal skeleton is prominent in profile from at least the third month
on. In both gorilla and chimpanzee it is visible early in fetal life
but soon vanishes. In the smaller Old World monkeys, and even
in the baboon Papio hamadryas, it is also present fetally.
Among the living primates this kind of nasal profile is a very
primitive feature found in the adult form in lemurs, tarsiers, most
of the South American monkeys, and the smaller and less spe-
cialized Old World monkeys.2 Man’s prominent nasal skeleton,
which serves as the roof of a resonance chamber useful in speech,
9 B. Kummer: “Untersuchungen iiber die Entwicklung der Schadelform des
Menschen und einiger Anthropoiden,” AEB, No. 3 (1953), pp. 1-44.
Bolk: op. cit.
1 de Beer: op. cit., p. 58, after Coupin.
2 See particularly the drawings of primate skulls in W. K. Gregory: Evolution
Emerging (New York: The Macmillan Company; 1951), Vol. II.
Differences in Postnatal Growth
171
is thus an ancient primate possession, part of the precatarrhine
complex. In so far as man retains it for a new purpose, he is less
catarrhine than the Old World monkeys and apes. As we shall see
later, some living human races are more catarrhine than others in
this respect. We cannot, therefore, call a flattish nasal skeleton
pedomorphic.
Differences in Postnatal Growth
As our study of fetal differences in primates has been push-
ing us steadily past the zero hour of parturition, let us consider
postnatal growth, to round out the picture. Among the insects
there is no problem. Cinderella-like, the transition from larval to
adult life is an abrupt one: what was at last view a hairy, crawling
grub suddenly takes wing as a beautiful butterfly, as large and
perfect as it will ever be. Among the mammals no such dramatic
transformation takes place. An attenuated postnatal growth
period is in many ways an open-air prolongation of fetal life,
lasting anywhere from a few weeks in some tiny rodents to twenty
years in elephants and men. It lasts three years in the prosimians,
seven in the Old World monkeys, nine in the gibbon, and eleven
in the anthropoid apes.
In this postnatal growth period most of the differences between
monkeys, apes, and men which in the womb were so elusive now
take shape and assume proper proportions. The colobus’s thumb,
barely present at birth, remains a button or shrinks from sight;
the gibbon’s forearm grows mightily. No sooner are the great apes’
milk teeth all in place than the huge permanent teeth follow,
crowding them out; and the jaw early assumes its massive form.
Man s teeth pursue a more leisurely sequence, waiting for the
brain to reach nearly adult size before the permanent set begins
to erupt, at the age of five or six.
In the monkeys and apes the sutures of the skull close not long
after birth. This closing does not halt the brain’s growth, but it
signifies that the brain has itself ceased growing. In man the su-
tures do not close until much later, around the age of thirty.
When, by some genetic accident, the sutures close early in man,
the result can be a microcephalic idiot.
i72 Man’s Place among the Primates
Probably the most human characteristic of man’s development
is not so much his posture or his brain as the fact that from birth
to belated maturity it takes six times as many calories of food
per kilogram of adult weight to build a man as to nurture any
ordinary mammal to adulthood.3 Man, then, is the most expensive
of all animals to rear. Hence, the need of special techniques to
obtain extra calories must have placed an early premium, and a
survival value, on culture, which in turn depends on superior loco-
motion and brain power. The effects on language, the family,
and technology are self-evident.
Physiological Clues to Our Relationships with Other Primates
Comparative anatomy and the study of prenatal and
postnatal fetal growth are not the only approaches to taxonomic
relationships between primates. Scientists working with micro-
scopes and test tubes have disclosed a set of physiological similari-
ties and differences which are equally valuable, and usually quite
technical. New information is becoming available so rapidly that
only a specialist can keep up with it. I know, at the time of writing,
of nine physiological tests that relate man to other primates. The
first two concern the urine, the others the blood.
( 1 ) Purine Metabolism. Man resembles the apes and differs
from the other primates.4
8 M. Rubner: Das Problem der Lebensdauer und seiner Beziehung zum Wach-
stum (Munich and Berlin, 1908), after de Beer.
“Purine is a crystalline compound (C5H4N4), the parent of other compounds
of the uric acid group. All the primates in which purine metabolism has been
studied, except man and the apes, carry the oxidation of purine through the uric
acid stage to that of allontoin, which is excreted in the urine. In man and the
apes, the process stops with the production of uric acid, only about half of which
is passed in the urine, the other half circulating in the blood stream. This trait
is also found in one breed of dog, the Dalmatian; there it has been traced to the
recessive allele of a single gene. The circulation of uric acid in the blood stream
is said to have a stimulating effect, like that of caffein, on the physiology of the
brain. This explanation is flattering to humans and apes, but it is not supported
by the inclusion of the Dalmatian, whose intelligence does not surpass that of
other breeds.
W. L. Straus, Jr.: “Urine of Anthropoid Apes,” Science, Vol. 124 (1956),
P- 435-
W. C. O. Hill: Man's Ancestry (Springfield, Illinois: Charles C Thomas;
1955), P- 87.
Our Relationships with Other Primates 173
(2) The Rate of Excretion of Five Amino Acids in the Urine.
Adult human beings differ from apes, but human infants resemble
them in this trait. No data is available for other primates.5
( 3 ) ABO Blood Groups. Man resembles the apes, particularly
the chimpanzee, and differs from the macaques.6
S. M. Gartler, I. L. Firscheim, and T. Dobzhansky: “A Chromatographic In-
vestigation of Urinary Amino Acids in the Great Apes,” AJPA, Vol. 14, No. 1
(1956), pp. 41-58.
De W. Stetten, Jr.: “Gout and Metabolism,” SA, Vol. 198, No. 6 (1958)
pp. 73-81.
In the rate of excretion of five amino acids in the urine, all four apes, includ-
ing the gibbon, show marked differences from man. Man excretes much more
creatinine and histadine than apes, but apes exceed man in glutamic acid, aspartic
acid, and beta-alinine. Since human infants resemble adult apes in the excretory
ratio of glutamic acids, aspartic acid, and creatinine, it is conceivable that com-
pared to the apes man is gerontomorphic in these physiological processes. How-
ever, no reports are available to date on the excretion of these substances in other
primates, so we cannot use this test for a three-way comparison between man,
apes, and monkeys. But this study does serve to line man up in one taxonomic
camp and all four apes in another. Among ape genera, lesser differences may be
seen. Chimpanzees excrete more histadine than the others, gorillas more aspartic
and glutamic acids, and beta-alanine. In the largest nonhuman primate sample
studied, that of thirty-seven chimpanzees, individual variation was seen to be as
great as in man, which is also true of many other chimpanzee traits.
Gartler et al: op. cit.
The ABO blood groups are found in the Old World monkeys and apes as
well as in man. In man and all apes except the gorilla all the substances are
carried in the blood itself. In the gorilla the anti-A substance is carried in the
blood and the anti-B in the salivary glands. Both substances are carried in the
urine. In the Old World monkeys the substances are present only in body fluids
and other tissues, but not in the blood.
The chimpanzee has both groups O and A, and the A is neither Ai nor Aa, as
found in most human beings, but a third type called Ai., (Wiener & Gordon,
i960). This is also found in man, most frequently in Negroes, rarely in Cau-
casoids, and never in Mongoloids. The mountain gorilla has only group A; the low-
land gorilla only group B, and the B substance is different from its human
counterpart. Both orangs and gibbons have A, B, and AB. Among the monkeys
the rhesus ( Macaca rnulata ) has only B (Biichi, 1953), but the macaque of
Java ( Macaca irius) has A, B, and AB.
All thirteen chimpanzees studied for the secretor trait, which is associated
with the ABO system, were found to have it.
P. B. Candela, A. S. Wiener, and L. J. Goss: “New Observations on the Blood
Group Factors in Simiidi and Cercopithecidae,” Zoologica, Vol. 25, No. 4 (1940),
PP- 513-21-
E. C. Biichi: “A Rhesus Monkey with B Agglutinogen,” Nature, Vol. 172
(i953)» P- 873-
S. D. and L. J. Lawler: Human Blood Groups and Inheritance (Cambridge,
Mass.: Harvard University Press; 1957), p. 82.
A. S. Wiener and E. B. Gordon: ‘ The Blood Groups of Chimpanzees, ABO
Groups and MN Types,” AJPA, Vol. 18, No. 4 (i960), pp. 301-11.
1 74 Man’s Place among the Primates
(4) MN Blood Groups. Man resembles the chimpanzee more
closely than other primates.7
(5) Precipitin Test. Man is identical with the chimpanzee,
resembles Old World monkeys to a recognizable extent, and shows
no kinship to the New World monkeys or the prosimians.8
(6) Serum Albumin and Serum Gamma Globulin. Tests were
made on man, gibbon, macaque, mandrill, and marmoset. In these
characteristics man resembles the gibbon most, the macaque and
mandrill next ( and equally ) , and the marmoset least.9
7 Both the apes and the Old World monkeys have the M antigen of the MN
series, but in the three major groups, man, apes, and Old World monkeys, this
M substance varies in chemical composition. That closest to man is found in the
chimpanzee, which is also the only subhuman primate known to have the N
antigen.
However, when Wiener and Gordon tested thirteen chimpanzees for M and N,
they found all to be MN; not one was MM or NN. Such a distribution would be
impossible in man, because we inherit this trait in Mendelian fashion, and there-
fore only 50 per cent of any series could be MN. The chimpanzee type of MN
must therefore be inherited differently. Chemically only half the chimpanzees
tested had the human type of N substance. The others had a type specific for
chimpanzees.
Wiener and Gordon also tested the same chimpanzees, and one dead animal,
for an anti-U factor, related to the MN system. All fourteen lacked it. It is present
in the blood of all Caucasoids tested but absent in some Negroes. The same au-
thors also point out a similarity in the reaction to a certain Rhesus antigen (Rh-
Hr) between the bloods of chimpanzees and Negroes. They interpret these Negro-
chimpanzee similarities (Ai.2, anti-U, and Rh-Hr) as parallel mutations suited to
the African environment.
Lawler and Lawler: op. cit.
Weiner and Gordon: op. cit.
8 In the precipitin test a rabbit is immunized with small doses of human blood.
The serum from this rabbit’s blood is used, largely by the police, to identify
human blood. If the rabbit serum is mixed with human serum, a precipitate is
formed. Some cloudiness is produced if the rabbit serum is mixed with chimpan-
zee serum, less cloudiness if mixed with serum from the Old World monkeys, and
no reaction is seen at all when it is mixed with the sera of New World monkeys
or lemurs.
Hill: op. cit., p. 8.
P. Kramp: “Serologische Stammbauforschung,” Primatologia, Vol. 1 (Basel:
S. Karger: 1956), pp. 1015-34.
9 A refinement of the precipitin test involves the comparison of rabbit and
chick antisera with the serum albumin and serum gamma globulin of various mu-
tually related species. Goodman has compared, in this fashion, the gibbon, ma-
caque, mandrill marmoset, and man. His experiments have placed us closest, of
these four animals, to the gibbon. Next come the macaque and mandrill, about
equally close. Our kinship to the marmoset is traceable but most distant.
M. Goodman: “The Species Specificity of Proteins as Observed in the Wilson
Comparative Analyses Plates,” AN, Vol. 94, No. 875 (i960), pp. 184-6.
Our Relationships with Other Primates
175
(7) Hemoglobins, Haptoglobins, and Serum Transferrins.
Man and apes resemble each other, and the Old World monkeys
are radically different from both.1
(8) Whole Globulin Molecules. Man is almost indistinguish-
able from chimpanzee and gorilla. Of the other primates only the
orang was tested.2
(9) Gamma Globulin, Gm Group. Man is closer to the chim-
panzee than to the gibbon, Old World monkeys, or New World
monkeys.3
In these nine tests man’s biochemical relationship to apes was
explored. In six tests man was compared to Old World monkeys,
in three to New World monkeys, and in one to prosimians. Man
is shown to be closely related to the apes — more closely to the
chimpanzee than to either the gorilla or the orang. The gibbon is
more distant from man than the other apes are. Man’s relation-
ships to the other primates are even more distant.
Until further data is available, we may consider the order of
1 In a paper delivered before the American Association of Physical Anthro-
pologists on May 12, i960, J. Buettner-Janusch reported on work in progress on
Hemoglobins, Haptoglobins, and Serum Transferrins of a Number of Old World
Primates. Apparently serum haptoglobins are the same in all primates studied, in
that there are three types identical in each population, including prosimians: two
homozygotes and a heterozygote. However, differences are found in the serum-
transparent beta globulins, which are governed by eight or nine alleles. The beta
globulins difFer from species to species, and even within bands of a single species
(baboons). The pattern seen in the Old World monkeys is radically different
from that in apes and man. At the time of writing, Buettner-Janusch is engaged
in extensive research on this and similar physiological comparisons between pri-
mates. As the beta-globulin test discloses differences among populations within
a species, it should be useful for racial studies in man.
2 In a paper delivered before the American Chemical Society on September 18,
i960, Emile Zuckerkandl stated that the patterns of whole hemoglobin molecules
differ among animal species in accordance with their evolutionary relationships.
“The hemoglobin patterns of man and eleven adult animals were analyzed, in-
cluding the gorilla, chimpanzee, orangutan. . . . ‘The . . . technique shows
that man’s hemoglobin is almost indistinguishable from that of the gorilla and
chimpanzee,’ Dr. Zuckerkandl said.” ( The New York Times, September 18, i960.)
3 S. H. Boyer and W. J. Young examined the gamma globulin (Gm) of 24
chimpanzees, 2 gibbons, 25 baboons, 2 rhesus monkeys, 2 spider monkeys, 1 red
(?) monkey, 4 cows, and 5 mongrel dogs. Only the chimpanzee serum inhibited
any of several Gm type reactions. Chimpanzee sera, like those of man, are poly-
morphic, and in both species the GM phenotype is not associated with gamma
globulin concentration. In these three respects the chimpanzee of all animals stud-
ied is closest to man.
S. H. Boyer and W. J. Young: “Gamma Globulin (Gm Group) Heterogeneity
in Chimpanzees,” Science, Vol. 133, No. 3452 (1961), pp. 583-4.
176 Man’s Place among the Primates
resemblance between man and the pongids, in this characteristic,
to be: chimpanzee, gorilla, orang. Man is closer to the gibbon
that to the Old World monkeys, and closer to the latter than to
either the New World monkeys or the prosimians. For what it is
worth, these tests relate us more closely to the African pongids
than to any other primates yet tested.
Parasites and Primates
Still another test of man’s kinship to his fellow primates
involves mutual parasites, internal and external. Chimpanzees can
serve as hosts for malaria and can be given syphilis in a mild
form. This disease can also be given to baboons; it is even milder
in them and soon disappears. The chimpanzee can also be given
yaws. In Ruanda-Urundi the mountain gorilla can have an in-
testinal parasite in common with man. These comparisons are
suggestive, but their validity is weakened by the versatility of
disease organisms: trichinosis, for example, may pass from pigs to
bears.
However, external parasites are different. Owing to an extreme
biochemical specialization, a particular kind of louse, for example,
can live only on its habitual host or on another genetically very
close. The application of this principle to birds led to the discovery
that flamingoes, despite their long legs and specialized beaks,
are really ducks. Pursuing this line of evidence, we discover that
body lice of the genus Pediculus exist on man and the chim-
panzee. The Old World monkeys are infested with lice of another
genus, Pedicinus, and the New World monkeys, lorises, and lemurs
have one other genus each.4 Aside from Pediculus hurmnus on his
head and body, man supports a louse of another genus, Phthirus
pubis, in the neighborhood of his genitals, and the gorilla has a
louse of the same genus but of a different species, P. gorillae,
around and about his private parts. No other species of Phthirus
has been found.5
4 Hill: op. cit., pp. 8-10.
5 H. Levene and T. Dobzhansky: “Possible Genetic Difference. . . .”
R. R. Gates: Human Genetics (New York: The Macmillan Co.; 1946), Vol. 2,
pp. 1419-21.
The Comparison of Primate Chromosomes
177
The Comparison of Primate Chromosomes
The study of chromosomes is a relatively new tool of taxon-
omy, and a particularly useful one. So far as we know, chromo-
somes are not influenced by environment. Also, in a few species,
it has been possible to equate chromosome micro-anatomy with
the gross anatomy and functions of the whole organism.
Geneticists recognize two principal kinds of cells, haploid and
diploid. In a haploid cell there is only one set of chromosomes.
Only sperm cells and unfertilized egg cells are haploid. Diploid
cells have two sets of paired chromosomes. Fertilized egg cells and
autosomal cells — the ordinary cells of the body — are normally
diploid. The diploid figure is commonly used to indicate the num-
ber of chromosomes in the cells of an animal.
The number of chromosomes per nucleus in each species is
virtually constant. In normal human beings only about 1 per cent
of the diploid cells vary from the number 46." The numbers 45,
47, and 48 occur principally in hermaphrodites, persons con-
genitally lacking sexual parts, and others suffering from certain
hereditary diseases. These deviations are principally concerned
with the sex chromosomes. Similar individual variations have also
been observed in other primates.7
With these exceptions, the number of chromosomes is constant
within a species and often so within related species. Unless it is
the same, animals cannot produce healthy, fertile offspring. If two
or more supposed subspecies can be shown to have different
normal chromosome counts, then they are separate species. For
example, in the Brown lemur, Lemur fulvus, counts of 60, 52,
and 48 chromosomes have been made on three populations classi-
fied as subspecies ( see Table 2 ) .
Within the order of primates, counts made to date range from
72 to 34. We cannot assign segments of this range to special
families or subfamilies because each family or subfamily so far
fiJ- H. Tjio and T. T. Puck: “The Somatic Chromosomes of Man,” PNAS,
Vol. 44, No. 12. (1958).
7 E. H. Y. Chu and B. A. Swomley: “Chromosomes of Lemurine Lemurs,”
Science, Vol. 137, No. 3468 (i960).
178
Mans Place among the Primates
studied in which more than one or two species are represented
shows a wide range of its own. As is true among other kinds of
animals, within each subfamily the chromosome count is high-
est in the simplest, most generalized species, and lowest in those
most specialized. Man, with 46 chromosomes, lies a little below
the middle of the primate range. Of the 72 species studied, 35
have more chromosomes than man, 8 have the same number,
and 29 have fewer. Man has the same number as certain species
of lemurs, marmosets, and cebus monkeys, to which he is not par-
ticularly related; only the number 42 is as common as 46. On the
basis of the chromosome count it cannot be said that man is
closer to the chimpanzee, gorilla, and orang, who have 48 each,
than to the macaques and baboons, who have 42.
Raw numbers of chromosomes cannot be used on a simple, lin-
ear scale to indicate taxonomic relationships because chromo-
somes tend to combine as the animals become increasingly spe-
cialized. The individual chromosomes in a single cell vary greatly
in length. In man the autosomal chromosomes (all but the sex
chromosomes X and Y) vary from 1.8 to 9.6 per cent of their com-
bined length, or more than five to one. The total length of all the
chromosomes in a cell seems to be a more useful figure than their
number.
Chromosomes also vary in the position of their centromeres. A
centromere is a specialized segment of a chromosome which acts
both as an adhesive and as a repellent. It is the junction point of
the two strands of which the chromosome is composed at certain
stages. When the chromosome splits in cell division the centro-
meres act as the foci of separation.3
If the centromere is located more or less in the middle of a
chromosome, that chromosome is called metacentric. If it comes
at about the three-quarter point, so that the arms on the two
ends are of unequal length, that chromosome is subterminal. If
the centromere is set at the very end of the chromosome, the latter
is telocentric ( or acrocentric ) .8 9
8 For details, see any standard textbook on elementary genetics.
9 The thresholds between these categories seem to be arbitrary. Tjio and Puck
have invented a ratio by which the categories can be standardized — the length of
the long arm divided by that of the short arm. The figures range from 1.08 to 10.5.
The Comparison of Primate Chromosomes 1 79
Geneticists have devised a means of illustrating the relative
sizes and shapes of the chromosomes for any species. They line up
one each of the autosomal chromosomes according to size and
relative arm length, and also show both sex chromosomes. Such a
chart is called a karyotype ( see plate XXI ) . In the human karyo-
type none of the autosomal chromosomes are telocentric. The
male sex chromosome Y may or may not have a very short second
arm. Two chromosomes, Numbers 18 and 21, have curious-looking
antennae.
At any rate, two telocentric chromosomes can combine into one
metacentric unit by joining end to end. By this process the num-
ber of chromosomes can be reduced in one of several closely re-
lated species. Thus, for example, the goat ( Capra sp .) has 60
paired chromosomes, all telocentric. The sheep ( Ovis sp.) has 54,
of which 48 are telocentric and 6 metacentric. Apparently 12
telocentric chromosomes of their common ancestor became fused
into 6 metacentric ones to produce the sheep.1 Nevertheless, these
two animals aie so similar anatomically that a mammalian anato-
mist can hardly tell their bones apart, or even their teeth. Yet the
two genera cannot interbreed. Any good human anatomist can
tell the skeletons of human races apart; yet all human beings are
members of a single species and can interbreed. If in our survey
of the primate chromosome patterns we find two genera of mon-
keys or apes with different chromosome numbers, we must seek
further data before deciding on degrees of affinity.
Bender and Mettler suggest that the original number of chromo-
somes for the primates was about 70 (presently the maximum is
72), all being small telocentrics, and that this number has been
reduced by combination.
So far chromosomes are no more useful taxonomically, in the
sense of indicating genetic affinity, than a host of other charac-
teristics, but they may become so. Fruit-fly specialists have been
able to assign special functions to individual segments of individ-
The authors arrange these in four categories: median = o-i. 30; submedian =
1-31-3-13; subterminal = 3.13-10.50. According to this system, man has 6 median,
22 submedian, and 16 subterminal pairs. None are terminal.
1 S. Makino: “The Chromosome Complexes in Goat ( Capra hircus ) and Sheep
(Ovis aries), and their Relationship,” Chromosome Studies in Domestic Mammals,
II, Cytologia, Vol. 13, No. 1 ( 1943), pp. 39-54.
i8o
Man’s Place among the Primates
TABLE 2
NUMBERS OF CHROMOSOMES AMONG
THE PRIMATES*
Diploid, Chromosome Numbers
Subfamily Species
Source
Common Name
Number
Tupaiinae
Urogale everetti
D
Philippine tree shrew
26 (?)
Lemurinae
Lemur mongoz
C&S
Mongoose lemur
60
Lemur fulvus rufus
C&S
Brown lemur
60
Lemur fulvus sp. nov.
C&S
Brown lemur
52
Lemur fulvus fulvus
C&S
Brown lemur
48
Lemur albifrons
C&S
Black lemur
60
Lemur catta
C&S
Ring-tailed lemur
56
Lemur variegatus
C&S
Ruffed lemur
48
Lemur variegatus ( subspecies ) C&S
Ruffed lemur
46
Lemur macaca
C&B
Black lemur
44
Hapalemur griseus olivaceus C&S
Grey gentle lemur
58
Hapalemur griseus griseus
C&S
Grey gentle lemur
54
Microcebus murinus
B&C
Miller’s mouse lemur
66
Lorisinae
Periodictius potto
C&B
Potto .
62
Nycticebus cougang
B&M
Slow loris
50
Galaginae
Galago crassicaudatus
C&B
Thick-tailed bush baby
62
Galago senegalensis
Mat
Lesser bush baby
38
Callithricinae
Callithrix chrysoleucos
C&B
Golden marmoset
46
Callithrix jaccus
Ch
Common marmoset
46 (?)
Leontocebus illigeri
B&Mf
Red-mantled tamarin
46
Callimiconinae
Callimico goeldii
C&B
Goeldi’s marmoset
48
Cebinae
Cebus sp.
P
Ringtail monkey
54
Aotus trivirgatus
* Symbol
C&B
Author and Title
Owl-faced monkey
54
YPY
C. H. Yeager, T. S. Painter and R. M. Yerkes: “The
Chromosomes of the Chimpanzee,” Science, Vol. 91,
No. 2351 (1940), pp. 74-5.
C&G
Chu and N. H. Giles: “A Study of Monkey Chromo-
some Components,” AJPA Proceedings, 1957, Ab-
stract 70, pp. 452-3.
B&M
M. A. Bender and L. E. Mettler: “Chromosome Studies
of Primates,” Science, Vol. 128, No. 3317 (1958), pp.
186-90.
T&P
Tjio and Puck: op. cit., pp. 1229-37.
YMFJ
Young, T. Merz, M. A. Ferguson-Smith, and A. W.
Johnston: “Chromosome Numbers of the Chimpan-
zee, Pan troglodytes ,” Science, Vol. 131, No. 3414
(1960), pp. 1672-3.
C&S
Chu and Swomley: op. cit., pp. 1925-6.
HFD
J. L. Hamerton,
H. P. Klinger,
M. Fracatto, L. Decarli, F. Nuzzo,
L. Hulliger, A. Taylor, and E. M.
The Comparison of Primate Chromosomes 181
TABLE 2
NUMBERS OF CHROMOSOMES AMONG
THE PRIMATES*
Diploid Chromosome Numbers
Subfamily Species
Source
Common Name
Number
Cebus apella
B&M
Cinnamon ringtail
54
Cebus capucinus
B&M
Capuchin ringtail
54
Pithecia pithecia
B&Mt
Saki
46 (?)
Cacajao rubicundus
B&Mt
Uakari
46 (?)
Alouatta seniculus
B&Mt
Red howler monkey
44 (?)
Callicebus cupreus
B&M
Red titi
46
Saimiri sciureus
B&M
Squirrel monkey
44
Ateles geoffroyi
B&M
Hooded spider monkey
34
A teles paniscus chamek
B&M
Black-faced spider monkey
34
Ateles belzebuth
B&M
Golden spider monkey
34
Ateles arachnoides
Ch
Woolly spider monkey
34 (?)
Lagothrix ubericola
B&C
Brown woolly monkey
62
Cercopithecinae
Cercopithecus I’Hoesti
C&B
l’Hoest’s guenon
72
Cercopithecus mona mona
B&M
Mona guenon
66
Cercopithecus mona denti
Tap
Guenon
66 (?)
Cercopithecus mona campbelli
C&G
Campbell’s guenon
66
Cercopithecus aethiops sabaeus
C&G
African green monkey
60
Cercopithecus aethiops tantalus
C&G
African white monkey
60
Cercopithecus diana
C&G
Diana monkey
60
Cercopithecus neglectus
Tap
De Brassa’s guenon
60 (?)
Cercopithecus nictitans
C&G
White or spot-nosed guenon
66
Cercopithecus cephus
Ch
Mustached guenon
54 (?)
Erythrocebus patas
C&G
Patas monkey
54
Cercocebus torquatus torquatus
B&M
Sooty mangabey
42
Cercocebus torquatus lunulatus
C&G
White-crowned mangabey
42
Cercocebus albigena
Tap
Grey-cheeked mangabey
42
Cercocebus galeritus
Tap
Crested mangabey
42(?)
Macaca mulata
D&H
Rhesus macaque
42
Macaca irius
C&B
Crab-eating macaque
42
Ch
Lang: “Somatic Chromosomes of the Gorilla,” Na-
ture, Vol. 192, No. 4799 (1961), pp. 225-8.
B. Chiarelli: “Chromosomes of the Orang-utan ( Pongo
B&C
VVomaeus)," Nature, Vol. 192, No. 4799 (1961), p. 285.
Bender and Chu, with permission.
C&B
"Chromosome Cytology and Evolution in Primates,”
P
Science, Vol. 133, No. 3462 (1961), pp. 1399-405.
References Cited in C&B
T. S. Painter: 1924.
S
P. I. Shiwago: 1939.
Mak
S. Makino: 1943.
D&H
C. D. Darlington and A. Haque: 1955.
Mat
R. Matthey: 1955.
R&S
K. H. Rothfels and L. Siminovitch: 1958.
D
O. Dodson, in personal communication to Chu and
T
Bender.
N. E. Tappan, in personal communication to Chu and
BAMf
Bender.
M. A. Bender and L. E. Mettler: Unpublished.
182
Man’s Place among the Primates
TABLE 2
NUMBERS OF CHROMOSOMES AMONG
THE PRIMATES*
Diploid Chromosome Numbers
Subfamily Species
Source
Common Name
Number
Macaca nemestrina
D&H
Pig-tailed macaque
42
Macaca cyclopis
Mak
Formosan macaque
42
Macaca sylvana
Ch
Barbary ape
42
Macaca assamensis
Ch
Assamese macaque
42
Macaca fuscata
Ch
Japanese macaque
42 (?)
Macaca sylenus
Ch
Lion-tailed macaque
42
Papio papio
D&H
Guinea baboon
42
Papio sphinx
B&M
Mandrill
42
Papio doguera
C&G
Olive baboon
42
Papio leucophaeus
B&C
Drill
42
Colobinae
Colobus ( polycomos )
B&C
Colobus monkey
44
Presbytis entellus
Mak
Langur
50
Hylobatinae
Hylobates lar
B&C
White-handed gibbon
44
Hylobates agilis
B&C
Agile gibbon
44
Hylobates hoolockii
C&B
Iloolock (gibbon)
44
Symphalangus syndactylus
B&C
Siamang
50
Ponginae
Gorilla gorilla
HFD
Gorilla
48
Pan troglodytes
YPY
Chimpanzee
48
Pongo pygmaeus
Ch
Orangutan
48
Homininae
Homo sapiens
T&P
Man
46
ual chromosomes. All that human geneticists have accomplished
in this line is to determine some of the traits that are carried on
the sex chromosomes X and Y. They know that others are carried
on the autosomal chromosomes, singly or in groups, but they can-
not say which autosomes carry which traits. But experimental
cytogenetics is advancing rapidly, thanks to the electron micro-
scope and the use of biopsy samples from live individuals, and it
may not be long before we know the function of each segment
of each chromosome in the development of the human organism.2
When that time comes, we may be able to draw karyotype charts
2 Investigations made with the electron microscope at Cold Spring Harbor have
shown that in cells of an animal’s body single chromosomes excrete RNA from
nucleus to cytoplasm. If we can discover which chromosomes are involved in dif-
ferent parts of the body, man may become as well known genetically as the fruit
fly. See Helen Gay: “Nuclear Control of the Cell,” SA, Vol. 202, No. 1 (i960),
pp. 126-36.
The Evidence of Behavior 183
of human subspecies, and the study of race in man will be on
firm ground.
The Evidence of Behavior
The one remaining category of evidence concerning man’s
relationship with the other primates is behavior, which taxono-
mists now consider as important a criterion of species as anatomy
and physiology. This decision was prompted by the fact that in
the course of natural selection animal populations are pruned for
their individual capacities for behavior. In addition, selective be-
havior in mating makes species possible.
In man behavior takes two forms, technological and social.
We can dismiss technology as a basis of comparison because man
alone has it. (Such minor activities as temporary nest-building
among the apes can be disregarded. )
Social behavior involves both nonsexual and sexual activities,
both of which are concerned with family structure. In most if
not all primate species, as among many other animals, the mother
shows anxiety about the safety of her newborn young. Juvenile
primates belonging to the same band play together as children
do, exercising their muscles, learning motor habits, and establish-
ing interpersonal relationships. Monkeys and apes groom each
other’s bodies and when night falls some may sleep together for
mutual warmth, companionship, and protection. Human beings
do all these things and in this they resemble all the other primates
in general, except the prosimians, rather than any one family,
genus, or species of monkey or ape.3
In sexual behavior, which forms another basis of group organi-
zation, man does not strictly follow the pattern of any other pri-
mate family, genus, or species. Man’s closest kin, the three great
apes, live in simple harems. More complex simian societies in
which two or more adult males tolerate one another’s presence are
found among South American monkeys, Old World leaf eaters,
and the terrestrial Old World monkeys such as the macaques and
baboons.
Although these three groups of monkeys, which are only re-
3C. H. Southwick: “Letter to Editors,” SA, Vol. 203, No. 6 (i960), p. 14.
1 84 Man’s Place among the Primates
motely related to man and to each other, have also achieved a
social order in which adult males can live peacefully together,
their patterns of social behavior differ from man’s in other re-
spects. The club type of sex life practiced by the howler monkeys
has no counterpart in normal human society, although something
similar turns up in houses of prostitution and at times of war.
Among the baboons, when an adult female begins to come into
heat she is first served by one or more eager, youthful males; only
when in full oestrual bloom, as ripe as a persimmon, does she
crawl to the old king, who then deigns to serve her. This behavior
pattern has certain human counterparts that need not be over-
interpreted, such as the bachelor’s house in many primitive socie-
ties, patterned adultery among the Tiwi, and the noctural activity
of Turkish sultans.
The primate closest to man in family life is the gibbon, to whom
man is less closely related in other respects, as far as we know,
than he is to the great apes. But when we consider the human
trait of solicitude on the part of the male parent toward the help-
less young, man’s closest counterpart is the male marmoset, who
carries his wife’s babies about and weans them with premasticated
food.
The kind of society man lives in, then, does not relate him to the
kin that is closest to him anatomically and physiologically. Man
has moved into new realms in locomotion and communication,
and has developed a pattern of behavior of his own which finds
its closest parallels in his more distant primate kin. This similarity
may be due to neoteny of endocrine origin, to higher intelligence
derived from competition among males for the largest harems and
among females for the most desirable males, and to recombina-
tions of genetic possibilities inherent in the primate gene struc-
ture. That we do not usually behave like apes does not mean that
we are not genetically close to them.
Among living peoples vast gaps separate the behavior patterns
of simple hunters and root diggers from those of sophisticated
urbanites and exurbanites. Yet the hunters belong to all five geo-
graphical races and the urbanites belong to at least three. Be-
havior in this sense is not a matter of race.
Among the subhuman primates the species noted for highest
The Evidence of Behavior 185
intelligence — and all above the prosimians are bright animals —
belong to several families that also include species of lesser wit,
as far as we can tell. In any colony of chimpanzees individual dif-
ferences of mental aptitude are profound. The genetic basis for
high intelligence has been acquired independently in different
taxonomic categories of primates. There is no evidence that the
most successful populations within several different human races
have not also become bright independently. If we believe they did,
the maze of human evolution can be straightened into a multiple-
laned highway.
This is as far as the pursuit of comparisons among living pri-
mates can take us. The next step is to discard the rich evidence of
flesh, fur, blood, lice, chromosomes, and conniving, and to follow
the bare bones of our ancestors and their relatives backward in
time to the moment when the primates first appeared.
8
6
K
THE FOSSIL RECORD FROM
LEMURS TO SWAMP APES
-|« -r- On the Scarcity of Primate Fossils
-Lio matter how carefully we compare the anatomy,
physiology, chromosomes, and behavior of the living primates, we
shall not, by these means alone, completely solve the secret of
their mutual relationships, or of our own descent, for each species
is the end product of its own evolution. No species is standing
still; not one is identical with its ancestors which lived in the early
days of mammalian differentiation. By the same token, compara-
tive embryology, although a valuable discipline, offers nothing
more than a succession of fetal forms, some of which may have
been omitted by neoteny. And zoogeography does not include
extinct species.
To leam the details of the ancestral journey of any species or
group of related species through the caverns of time there is no
substitute for the study of the records of paleontology. Only
through this specialized, fragmentarily documented, and all-
encompassing discipline can we hope to answer the questions:
Who are we? Whence do we come?
Were we sapient horses or snails, our task would be easy. Their
records have been worked out from A to Z. Unfortunately, how-
ever, we are primates, kin to an untidy, grimacing lot, and mem-
bers of an order whose ancestors chose the worst possible places
to live and and the worst possible way of living, in terms of the
preservation of skeletal material.
Plants equipped with chlorophyl turn carbon dioxide and sun-
light into sugar; animals eat the plants and one another; and bac-
teria break down the leftover tissues into simpler substances.
On the Scarcity of Primate Fossils 187
which repeat the cycle. Were this not so, the surface of the earth
would be stacked high with logs and bodies. As scavengers
abound, nearly all dead animals are transformed into the tissues
of new generations and death ensures the continuity of life.
Some parts of dead animals, however, resist decay. The hard-
est, most durable part of a vertebrate’s body (except for birds)
is its teeth. Much of paleontology therefore rests on dental com-
parisons, just as some kinds of archaeology rely heavily on pot-
sherds. This is fortunate, because the sizes, shapes, and structural
details of teeth are hereditary, independent of environmental in-
fluence except wear, and unaffected by growth changes once
erupted.
Even teeth, however, are hard to find in tropical forests where
the rainfall leaches away the topsoil and where the subsoil is acid.
The world’s great fossil beds are located in grasslands, and par-
ticularly in swampy terrain, where animals now and then get
mired, just as rhinoceros do today in muddy water holes during
the dry season. Unable to get out, they sink below the surface
of the mud and die, safe from predators, to be found millions of
years later embedded in sedimentary rock. This happens rarely
to primates. Most of them live in trees and never encounter mud.
Only the genera that live on the ground, like macaques and ba-
boons, frequent waterholes, and only their bones turn up in any
abundance. Our earliest known hominid predecessors also lived
in the open and quenched their thirst on the ground. Later, some
of them began to live in caves, where scraps of their bones turn
up in garbage heaps. Not more than seventy thousand years ago
did they begin to bury their dead. Only from the latest geological
period, therefore, are hominid bones at all frequent; anthropolo-
gists still have much less material to work with than do palentolo-
gists who trace the evolution of horses and rodents.
The Primate Time Scale 1
The primate fossil record covers the entire Cenozoic era. The
duration of this period is estimated at 63 million years on the
1 Geological time is officially divided into eras , periods , epochs , and ages. These
terms are often used loosely or interchangeably. However, they are defined as fol-
lows. All geological time consists of five eras, the Archaeozoic, the Proterozoic,
the Paleozoic, the Mesozoic, and the Cenozoic. Each era contains a number of
x 88 The Fossil Record from Lemurs to Swamp Apes
TABLE 3
THE CENOZOIC ERA IN MILLIONS OF YEARS
Duration
Ended X Millions
PERIODS
EPOCHS
Each
of Years Ago
Quaternary
Pleistocene and Recent
1
1
T
Pliocene
11
12
e
Miocene
13.7
25.7
r
Oligocene
8.3
34
t
Eocene
21
55
i
Paleocene
23
78
ary
basis of studies of the decay of uranium into lead, and by other
methods. It is divided as shown in Table 3. No division is made
periods. Those of the Mesozoic are Triassic, Jurassic, and Cretaceous; those of
the Cenozoic, Tertiary and Quaternary. Each period is divided into epochs. The
epochs of the Tertiary are Paleocene, Eocene, Oligocene, Miocene, and Pliocene;
those of the Quaternary are Pleistocene and Recent. The word age is used inde-
pendently of the other terms to designate the time span of a form of life or special
geological condition, thus: the Age of Fishes, the Age of Reptiles, the Age of Mam-
mals, and the Ice Age. Archaeologists and historians use some of these words in
special senses, e.g., the Stone Age, the Iron Age, the Hallstatt (Iron Age) Period,
and the Christian Era. The dates giving the divisions of the Cenozoic on Table 3
are based on J. L. Kulp: “Geologic Time Scale,” Science, Vol. 133, No. 3459
( 1961), pp. 1105-14.
Primate Paleontology as a Whole 189
in this chart between the Pleistocene and the so-called post-
Pleistocene, or Recent, which covers the time span elapsed since
the recession of the last glacial ice sheets in Europe and North
America at about 8,000 b.c. In other parts of the world it is more
difficult to determine.
Primate Paleontology as a Whole
During the Paleocene and Eocene the earth’s surface was
much smoother than it has been since, and tropical forests ex-
tended much farther poleward than they do today. Consequently
fossil primates are found in regions now uninhabitable for any
free-living primate except man. During the Oligocene the Alpo-
Himalayan and Rocky Mountain systems began to rise, and they
continued to do so during the Miocene. These new highlands
cooled off much of the poleward land of the Northern Hemi-
sphere which had formerly been suitable for habitation by ar-
boreal primates, isolating most of them in the three present-day
tropical faunal regions (Ethiopian, Oriental, and Neotropical).
During the Pliocene and the Pleistocene interglacials, however,
parts of southern and western Europe remained frost-free and
habitable by subhuman primates.
Unfortunately, the record of fossil primates is incomplete. More
investigation has been carried out in some countries than in others.
Moreover, in many areas whole epochs are completely un-
represented. In Africa south of the Sahara, a very accessible re-
gion where much work has been done and where hominid evolu-
tion may have gone on during the Pliocene, there are few known
deposits of that period. In the Dasht-i-Lut desert of southeastern
Iran, where Pliocene deposits are abundant, the ovenlike cli-
mate and general inaccessibility make paleonotological explora-
tion virtually impossible.
Certain elements of primate history can be explained by the rise
and fall of bridges between continental and off-shore land masses.
Madagascar was cut off from Africa in the Jurassic epoch of the
Mesozoic era, about 160 million years ago, when mammals were
just beginning to evolve. More than 100 million years later, dur-
ing the Eocene, the ancestors of the lemurs crossed to the island
from Africa by some unknown means, possibly by a temporary
190 The Fossil Record from Lemurs to Swamp Apes
land bridge. During the Miocene an incomplete bridge admitted
two fresh-water aquatic mammals, a dwarf hippopotamus and a
South African river pig, but no species arrived that could inter-
fere with the arboreal life of the lemurs until man appeared, some
2,000 years ago.
Africa and Eurasia were connected intermittently, mostly at
Suez as today. At one time or another these temporary bridges
allowed passage to all Old World primate forms. From the Cre-
taceous into the mid-Eocene, the Bering Strait was a land bridge
over which prosimians crossed from east to west; then it was bro-
ken until the Pleistocene, when man, along with many other land
mammals, crossed it.
During the Cretaceous and early Paleocene the Isthmus of
Panama connected North and South America, enabling prosimi-
ans to go south into the Neotropical forests, where they evolved
into the South American monkeys and proliferated mightily.
Meanwhile in North America the prosimians became extinct. Un-
til late in the Pliocene the isthmus remained under water. Then it
re-emerged, and it has been a land bridge ever since. After its rise
from the deep a number of dominant mammalian species went
south, extinguishing many of the hitherto sheltered local species;
but this invasion did not appreciably affect the primates, as it
included no competing arboreal forms. Moving in the opposite
direction, some of the South American monkeys ventured north
to Mexico, and men came down from the northern continent late
in the Pleistocene or even during the recent period. As Australia
had no Cenozoic land bridge, no primates came there until man
arrived, by island hopping, at some time late in the Pleistocene
when the Sunda and Sahul shelves were above water.
The Prosimian Proliferation
During the early Cenozoic — Paleocene to mid-Oligocene —
four orders of small mammals competed for a special ecological
lebensraum by developing, in some families, gnawing or chiseling
incisors like those of living beavers and squirrels. These were the
multituberculates, primates, rodents, and lagomorphs (hares
The Prosimian Proliferation
191
and rabbits). First to flourish, the multituberculates reached a
peak in the Paleocene but became extinct in the Eocene, possibly
because their incisors lacked dentine on the back with which to
keep a sharp edge, and their roots failed to continue growing
throughout life to replace wear, as the incisors of rodents do.
Next came the primates, then consisting exclusively of prosimi-
ans. They may have competed with the multituberculates and
helped bring about their downfall. Some fifty-five genera of early
Tertiary prosimians have been discovered — the better-known
are grouped in five families. Before the mid-Oligocene, three
families had become extinct. These were all chiselers and gnaw-
ers, probably forced out of action by competition with the rodents
and lagomorphs. The two surviving families, the Adapidae and
Anaptomorphidae, were nongnawers and owed their continued
existence to a lack of specialization. The only latter-day gnawing
primate is the aye-aye, who probably took up this habit later and
on his own.
The third and fourth chiseling orders, the rodents and lago-
morphs, were present from the Paleocene on but radiated only
after the decline of the gnawing prosimians. Their advantage
over the latter was physiological and behavioral rather than den-
tal. They are very fertile animals, with short pregnancies and
large litters. They are well adapted to life in deserts and cold
climates because they build nests and burrows and collect and
store food for lean seasons, and because some of them can live
without water and others hibernate. These adaptations gave them
dominance outside the tropical forests, where the nongnawing
prosimians were left to evolve after their own fashion. This in-
cluded long pregnancies, single births, and a long period of mat-
uration— features disadvantageous perhaps in the Paleocene and
even in the Oligocene, but essential to the eventual rise of man,
who builds houses, stores food, transports water, and proliferates
mightily by outwitting his competitors.
The Adapidae, found in both Europe and America, were un-
specialized lemurlike primates with small brains and the dental
formula - They were either the ancestors of living lemurs
*i*3
and lorises or their close kin, but no kin of man.
192 The Fossil Record from Lemurs to Swamp Apes
The Anaptomorphidae, however, with subfamilies in the Old
and New Worlds, were probably unspecialized tarsiers which had
branched off even earlier from the protolemur stem. From this
family all the living monkeys, apes, and man were probably de-
rived, before the ancestors of the living tarsiers had acquired
their specializations for hopping and nocturnal vision.
An important structural change that took place between the
protolemur and tarsius stages was a backward and sideward
shift of the line of stress on the skull which accommodates the
muscular pull of the jaws. In the lemurs, as in the shrews, much of
this stress is carried to the top of the head by way of the bony
framework between the eyes, which are consequently set far
apart and are not fully binocular. In the tarsiers this stress is
shifted mostly to the side of the face and head, outside the orbital
rims and behind the orbits. The eyes are then set closer together
and are fully binocular. These changes, which may be related to a
diminishing sense of smell and an increasingly better eyesight,
were carried over into both higher primate suborders, to a more
marked degree in the catarrhines than in the platyrrhines.
The Evolution of the Platyrrhines
From a North American anaptomorph the South American
monkeys evolved into four-footed limb crawlers, brachiators,
and tail-swingers ; and there we leave them. Their only bearing
on this story is that the same evolutionary parallelism occurs
in the differentiation of the Old World primates. None of them,
apparently, went down to the ground.
The Evolution of the Catarrhines
Meanwhile, the ancestors of the Old World monkeys, apes,
and men were evolving independently of the New World mon-
keys, apparently from one or more of the Old World anapto-
morphs, but exactly when and where we do not know. There is
The Evolution of the Catarrhines 193
even a slight doubt whether the Cercopithecoids and Hominoids
made the transition from prosimian to catarrhine in a single
evolutionary act, through a common ancestor, or whether the two
superfamilies independently crossed what paleontologists call an
adaptive threshold.
In Chapter 5 we discussed some considerable differences be-
tween the two superfamilies which do not concern locomotion or
posture. The Old World monkeys have double-disc placentas;
the apes and men, single-disc ones. Men and the apes me-
tabolize purine only partially; the monkeys completely.
In the apes and men, ABO blood-group substances are carried
in the blood; in the monkeys these substances are carried in
plasma and other media. The precipitin test demarcates the two
groups sharply, as do the genera of body lice with which each is
infested. As far as we know, chromosome counts likewise dif-
ferentiate the two groups. As long ago as 1945 Simpson divided
the New World monkeys, Old World monkeys, and the apes and
men into three superfamilies, the Ceboidea, Cercopithecoidea, and
Hominoidea, implying the separate descent of each from a lower
primate grade.2
Recent discoveries in paleontology have supported this position.
Until a few years ago two very small and ancient fossil mandibles
were believed to provide a common catarrhine link between the
Old World monkeys and the Hominoids. One was Amphipithecus,
found in an Upper Eocene deposit in Burma; 3 the other was
Pai apithecus, from the Lower Oligocene beds of the Fayum in
Egypt. Now both of these have been discredited.6 Not only do we
2G. G. Simpson: “The Principles of Classification . . . ,” p. 184.
3 E. H. Colbert: “A New Primate from the Upper Eocene Pondaung Forma-
tion in Burma,” AMN, No. 951 (1937).
4 W. E. LeG. Clark: “New Paleontological Evidence Bearing on the Evolution
of the Hominoidea,” QJGS, Vol. 105, Part 2 (1949), pp. 38.
5J. Hiirzeler: Oreopithecus bambolii Gervais, A Preliminary Report,” VNGB,
Vol. 69, No. 1 (1958), pp. 1—48, especially 32—3.
C. L. Gazin: “A Review of the Middle and Upper Eocene Primates of North
America, ’ SMC, Vol. 136, No. 1 (1958), pp. 1-112.
Clark: The Antecedents of Man (Chicago: Quadrangle Books; i960).
Hiirzeler rejects both Parapithecus and Amphipithecus and also two other du-
bious Fayum specimens, Apidium and Moeropithecus. Gazin casts doubt on Am-
phipithecus by inference, calling it “. . . a possible primate with three molars
194 The Fossil Record from Lemurs to Swamp Apes
lack a common ancestral catarrhine, but we have no fossil
Cercopithecidae older than the Lower Miocene, at which time
genuine Old World monkeys appear in East Africa. A small
frontal bone from the Lower Oligocene beds of the Fayum has
recently been identified as that of a primate, probably a catar-
rhine,6 but this does not solve the problem because we do not
know what kind of a catarrhine it was. In its general configura-
tion it resembles an ape rather than a monkey, but that is in-
conclusive.
The genuine Old World monkey found in the Lower Miocene of
East Africa is MesopithecusJ Sometime later, probably no more
then ten million years, the same genus is found in Greece,
Czechoslovakia, and Iran in a period known as the Pontian,
which the French call Late Miocene and the British and Germans
Early Pliocene. The best site is at Pikermi, in Greece, where com-
plete skeletons have been found. This animal is listed in the sub-
from the Eocene of Burma.” Clark expresses caution about the status of Parapi-
thecus, and W. L. Straus, Jr., who saw an enlarged photograph of Parapithecus
after it had been freshly cleaned (courtesy of Hiirzeler), states that the tooth
originally called a canine is a premolar, making the dental formula l : l : 3 : 3.
Straus feels that if Parapithecus was a primate at all it was an aberrant tarsier.
6 E. L. Simons: “An Anthropoid Frontal Bone from the Fayum Oligocene of
Egypt: the Oldest Skull Fragment of a Higher Primate,” AMN, No. 1976 ( 1959).
7 j. Piveteau: Les Primates et I’Homme, Traite de Paleontologie Humaine,
(Paris: Masson et Cie; 1957), pp. 135-43. This reference covers the rest of this
section, unless otherwise specified.
The Evolution of the Catarrhines 195
family Colobinae because its skull, jaws, and teeth resemble those
of the living leaf-eating monkeys of Africa and Asia. Among other
resemblances, the lower third molar has a third loph, as is true
of living colobines; and the lower molars and premolars are par-
ticularly worn on their outer edges whereas the corresponding
upper teeth are worn on the inner edges. This pattern of wear
favors the mastication of leaves, i.e., browsing.
The rest of the skeleton, however, is less specialized than that
of the living colobines and closer in form to those of macaques
and baboons. The femur is longer than the humerus, as is true of
the ground-living genera; the hand bones are longer than those
of leaf eaters and shorter than those of macaques and baboons.
The thumb is unreduced. The ischium has an enlarged, corru-
gated area suited for large ischial callosities like those of macaques
and baboons. In Kenya, where Mesopithecus remains are earliest,
they were part of a grassland fauna.8
On the whole, it looks as if this animal was close to the common
ancestor of the colobines and the ground-living Cercopithecines,
but had passed the taxonomic frontier into the colobine camp in
terms of diet while it still had some distance to go in locomotor
adaptation. It persisted in France until the Early Pleistocene,
and has turned up in the Pliocene deposits of the Siwalik Hills of
northern India and in the Pleistocene of Madras. Its route be-
tween Africa and India seems to have followed the Nile to the
eastern Mediterranean, the Fertile Crescent, and Iran, bypassing
southern Arabia.
To date, there are no true Cercopithecines available in the fos-
sil record before the Pliocene, when macaques ( Macaca prison
and others) appear in France, Holland, and Italy. Macaques are
also found in the Pleistocene of Europe, Indochina, and China.
The baboons first appear in the Late Pliocene or Early Pleistocene
of East Africa.
For one reason or another, including the fact that certain fossil
specimens just haven’t been uncovered, the fossil record of the
Old World monkeys is shorter than that of the Hominoids. More-
s B. Patterson: “The Geological History of Non-Hominid Primates in the Old
World, HB, Vol. 26, No. 3 (i954),pp. 191-219.
196 The Fossil Record from Lemurs to Swamp Apes
over, the Old World monkeys did not acquire their present spe-
cializations any earlier than apes and men did.
The Gibbon Line
The earliest known specimens of the Hominoids are rep-
resentatives of the least highly evolved of the three living families,
the Hylobatidae, or gibbons. At least three excellent sets of speci-
mens shed light on their evolution. The earliest is Propliopithecus,
a small, nearly complete mandible unearthed, like that of Para-
pithecus, in the Oligocene beds of the Fayum. This bone is only
two thirds the size of that of a living lar gibbon; if the body was
in proportion to the jaws the animal weighed only 7 to 10 pounds.
This places it within the weight range of a house cat.
Seen from above, the mandible is essentially V-shaped like those
of prosimians. Seen from the side, its ascending rami (the paired
branches behind the teeth which articulate with the skull) are
higher than those of a modern gibbon, suggesting a higher face.
Yet the mandible is shorter anteroposteriorly (from ear to lips)
and the chin line seems to have been nearly vertical. The molars
had five cusps and the premolars two. Although broken off, the
canines seem to have been of normal length for a gibbon; the
first premolar is not fully sectorial, that is, with a shearing edge;
and the incisors are missing. As the bone of the mandible is mas-
sive in proportion to the size of its teeth, the animal must have
had a powerful bite. Schlosser,9 who described it, considered it a
basal form of the gibbon family, but there is no apparent reason
why it could not also have been ancestral to the other apes, al-
though its relationship to the hominids is harder to see.
Moving on to the Lower Miocene, an animal named Limnopi-
thecus undoubtedly a gibbon, has been found on Rusinga Is-
land in Lake Victoria in Kenya. Several jaw fragments, teeth, and
9M. Schlosser: Oligoziine Landsaugetiere aus dem Fayum, BPGO, Vol. 51
(1911), pp. 51-167 (after Gregory, 1951).
1 Clark and L. S. B. Leakey: Fossil Hominids of East Africa, BMFM, Se-
ries 1, 1951.
Clark and D. P. Thomas: Associated Jaws and Limb Bones of Limnopithecus
macinnesi, BMFM, Series 3, 1951.
The Gibbon Line
197
limb bones have been recovered. The canines are shorter than
those of living gibbons and the anterior lower premolars still less
specialized for shearing. The incisors are a little smaller, and
the jawbones more robust. The limb skeleton, much of which is
preserved, is in some ways intermediate between those of liv-
ing Old World monkeys and gibbons. The arms are shorter than
those of living gibbons, and the whole rotating apparatus of the
shoulder girdle and elbow is only partly developed for brachia-
tion. This gibbon had only begun his career as a trapeze artist and
still lacked much of the necessary equipment. In fact, his legs
were quite strongly developed, and anatomical details of his foot
suggest more jumping or walking than modern gibbons indulge in.
In the Middle and Upper Miocene of Europe another ancestral
gibbon turns up. Pliopithecus 2 may in some future taxonomic re-
shuffling come to be included in the same genus as Limnopithe-
cus. Limnopithecus is represented by only a few teeth and scraps
of bone, whereas almost every bone in Pliopithecus’ s body below
the mandible has been recovered, as well as portions of the skull.
The dentition of Pliopithecus is similar to that of Limnopithe-
cus, and the mandible, like that of the living siamang — an animal
of the same body size — has a trace of a simian shelf, a bony strut
across the inside of the jaw at chin level. This mechanical prop
probably has no phylogenetic significance because it appears
when needed in big-toothed and heavy-jawed primates, and dis-
appears when no longer useful. A trace of prosimian influence
is seen in the form of the jaw, which is still V-shaped in the pos-
terior portion. The modern gibbon has lost this. Also, in what is
left of the facial skeleton, the interorbital distance is greater and
the nasal region wider than in modern gibbons, again a prosim-
ian relic.
The body of this animal is gibbonlike in details of the pelvis
and vertebrae. The sternum is broad and flat, as in brachiators
and man, and the clavicle (collarbone) is S-shaped as in orang-
2 H. Zapfe: “Die Pliopithecus-Funde aus der Spaltenfiillung von Neudorf an
der March (Czechoslovakia),” VGBV, Sonderheft C. (1952). Reprinted in Year-
book of Physical Anthropology (New York, 1951), pp. 55-g.
Zapfe: “Results of Research on the Skeleton of Pliopithecus ( Epipliopithecus )
vindobonensis.” Paper read at the Annual Meeting of the Am. Assn, of Phys. Anth.,
Cambridge, Mass., April 12, 1958.
198 The Fossil Record from Lemurs to Swamp Apes
utans and chimpanzees, rather than simply bowed, as in modem
gibbons. The hind legs are as long as a siamang’s, but the arms
are only 60 per cent as long as those of the living ape. The
shoulder and elbow joints are not as swivel-formed as in the mod-
ern animal, and both the radius and ulna articulate with the
carpal bones in prosimian and monkey fashion.
The tarsal bones of the feet are longer and the metatarsals and
toe bones shorter than in the siamang; and the same proportions
are found in the hand bones. In short, Pliopithecus was not yet a
full-time brachiator, nor was he altogether an arboreal quad-
ruped. He was a small ape that seems to have come down from the
trees, partly adapted himself to a quadrupedal life on the ground,
and was only beginning to become adapted to a renewed arboreal
life as a brachiator. In those anatomical details in which his limbs
differed from those of modern gibbons, according to Zapfe, he re-
sembled the prosimians as much as or more than the living Cer-
copithecid monkeys. The significance of this is uncertain because
Miocene Cercopithecids may have also resembled prosimians in
these features.
Limnopithecus and Pliopithecus establish the presence of an-
cestral gibbons in the Miocene of Europe and Africa; compar-
able remains of equal age have not been found in Asia. By the
Miocene the gibbons had already branched off from the stock of
the still-missing common ancestor of all the catarrhines, if indeed
such a common ancestor ever existed. By that time they had also
begun to become distinct from the ancestors of the Pongidae, or
great apes, and possibly also of the Hominidae, including Homo.
The Ancestors of the Three Living Great Apes
During the Miocene, the period in which the ancestors of the
gibbons appeared, the ancestors of the chimpanzees and gorillas
also made their first recognizable bow, and from the same general
stock. Although the ancestors of the orangutans must have ex-
isted at the same time, their bones have not yet been found or
identified. Simpson, in 1945, 3 and Fiedler, in 1956, 4 classified all
3 Simpson: op. cit.
4W. Fiedler: “Ubersicht iiber das System. . . .”
Proconsul
199
the ancestral great apes in one subfamily, Dryopithecinae, and
called their living descendants Ponginae. Some of the Dryopithe-
cinae became extinct by evolving into new forms, others by simply
dying out, apparently without issue. Thus the terms Dryopithe-
cinae and Ponginae represent evolutionary grades rather than
individual lines of descent, because both the chimpanzee and the
gorilla, in Africa, and the orangutan in Asia, must have evolved
independently from Dryopithecine ancestors.
Proconsul
The African Dryopithecines belong to a single genus, Proconsul,
named after a chimpanzee called Consul who lived in the London
zoo. Proconsul was found on the same fossil-rich island of Rusinga
as Limnopithecus. During the Miocene this part of the island was
apparently a forest, and the land below the trees was seasonally
flooded. This ape has three species and sizes: P. africanus, little
bigger than a gibbon; P. nyanzae, chimpanzee-sized; and P. ma-
jor, as big as a gorilla.5
The three are much alike except for size and the consequent
differences in proportions. As many more remains of P. africanus
have been found than of the other two, it has received the most
attention. Being the smallest and lightest, it was also, we believe,
the best brachiator of the three.
Unique among primate fossils of this age is an almost complete
adult female skull of P. africanus. As might be expected of a fe-
male of the smallest of the three species, it has a more or less
globular brain case, a forehead running at an angle of about 55 0
to the eye-ear plane ( a line drawn from the top of the ear hole
to the lower border of the orbit, which places the skull in its
normal operating position), no brow ridges, and an upper facial
skeleton set at the same angle as the forehead. The forehead slope
of a female gorilla is about 35°, and a modern human female
who happens to have a vertical forehad has, of course, a slope of
90 0 • Many women have less of a slope than that.
5 Clark and Leakey: The Miocene Hominidae of East Africa, BMFM, No. 1
(i95i)-
J. R. Napier and P. R. Davis: The Fore-limb Skeleton and Associated Remains
of Proconsul Africanus, BMFM, No. 16, 1959.
200
The Fossil Record from Lemurs to Swamp Apes
Although the cranial capacity cannot be measured accurately,
it probably lies somewhere between that of a gibbon ( ca. 100 cc. )
and that of a chimpanzee (ca. 400 cc.), which fits its body size.
As far as the endocranial cast can be read, it shows a frontal area
more monkeylike than apelike. In other words, the brains of these
Kig. 14 Proconsul africanus. Note that the skull of Proconsul africunus retains
several prosimian or platyrrhine-like features, including the relative position of the
eye sockets and nasal skeleton and the V-shaped lower border of the nasal
opening. The canines do not interlock very far; the bite of the unworn teeth is
intermediate between those of man and apes. ( Drawings after LeGros Clark and
Leakey, 1951.)
fossil apes had not evolved to the modern pongid level by the
Miocene.
As in some primitive men but not in living apes, the orbits are
low and broad, and the distance between them is great. Seen
from above, the orbits seem to face to the side more than they do
in living apes or even in most living Old World monkeys; in this
Proconsul
201
respect Proconsul resembles some of the lower primates which
lack full stereoscopic vision. The lower border of the nasal open-
ing is V-shaped, as in some Old World monkeys. In apes and
men it is a horizontal line.
A more modern feature is that the plane of the occlusal border
of the teeth, where uppers meet lowers when the jaws are closed,
as seen from the side, is parallel to the eye-ear plane, as in most
living apes (in the orangutan it actually slopes upward and fore-
ward ) and in most men. This sharply distinguishes them from the
living ground-monkeys, such as the baboon, whose tooth lines
slope downward and forward at a 40° angle.
The lower jaw is not massive and lacks a simian shelf. Its sides
are convergent, as among some lower primates; it does not have
parallel sides, as among the living apes, nor is it U-shaped as in
men. That the musculature of the jaw was relatively light is shown
by a wide separation of the temporal lines on the parietal bones,
by a medium development of the attachment areas for the mas-
seter muscles, and by a rather frail zygomatic arch.
The teeth, too, are not as impressive as those of modern apes.
The incisors are man-sized, the canine larger but moderate for
an ape, and in the maxillary bone of the face, above the root of
the canine, there is a small depression known as a canine fossa,
which is present in man but not in living apes. As in apes but not
in men, the first lower premolar is sectorial, that is, it has a shear-
ing edge. From numbers 1 to 3 the molars increase in size, and
number 1 is quite small. In living apes the second is largest and
the third smallest. The teeth of the other two species are similar
but larger, and the jaw of P. major is gorilloid in its massiveness.
Limb bones of all three species have been found and described,
but, as with the skull, the limb bones of P. africanus are the most
nearly complete. Napier and Davis have described an almost com-
plete left forelimb. Because of brachiation, the forelimb is criti-
cal in the identification of a pongid. In the humerus, radius,
ulna, carpal bones, metacarpals, and phalanges, this animal
showed some features reminiscent of the quadrupedal arboreal
primates and other features unique to brachiators. No features
unique to terrestrial quadrupeds like the macaques and baboons
were found. On the other hand, some quadrupedal traits in the
202
The Fossil Record from Lemurs to Swamp Apes
forelimb may be shared with arboreal and terrestrial forms; only
the brachiators are set apart. As P. africanus was only a part-time
or halfway brachiator, we cannot be sure that his ancestors, be-
fore beginning to brachiate, had not spent some time both in the
trees and on the ground.
The hand bones of ground-living monkeys are specialized for
walking and digging, as witness their long metacarpals and short
phalanges. The hand bones of apes, being specialized for brachia-
tion, have long metacarpals, long phalanges, and short thumbs.
The hand bones of P. africanus occupy an intermediate position,
one which indicates no complete form of specialization. He had
long phalanges, like an ape, but he also had a fairly long thumb,
like both arboreal and terrestrial monkeys, and man. The wrist
bones (carpals) were like those of the arboreal quadrupeds
rather than like either of the other forms, and the ulna met the
carpal bones as in quadrupedal primates.
As for the lower extremity, the upper end of the femur is ape-
like in general architecture, and the angle of head to shaft sug-
gests a carrying angle, as in apes and men but not in monkeys
either tree-borne or grounded. Napier and Davis have described
a nearly complete foot of P. africanus, which they find to be
largely apelike in the shortness of the tarsals, the proportions of
metatarsals to phalanges, and the divergence of the great toe.
The splendid work done on the Miocene primates of East Africa
by Leakey and his associates, among others, has given us a likely
ancestor for the chimpanzees and gorillas, and possibly also one
for man.
Dryopithecus in Europe and Asia
While the ancestors of the chimpanzee and gorilla were evolv-
ing in Africa, a much larger number of Dryopithecine genera were
similarly engaged in Europe and Asia. The entire subfamily is
named after a mandible found in France in 1856 by Lartet, and
dated in the mid-Miocene. He called it Dryopithecus fontani.
The genus Dryopithecus has since been found in other parts of
Europe, where some species persisted into the Pliocene.6 With two
6 Piveteau: op. cit., pp. 197-206.
R amapithecus, a Possible Ancestor of the Hominids 203
exceptions the specimens are limited to teeth and pieces of jaws.
One humeral shaft from France has been uncertainly labelled
Dryopithecus fontaniJ As both ends of it are missing, this hume-
rus tells us little. One complete femur, Paidopitliex, found in Ger-
many and formerly attributed to Dryopithecus, is listed under
the gibbons in Simpson’s compilation.8 Both bones are gibbon-
sized.
Recently two sets of teeth attributed to Dryopithecus keiyua-
nensis have been found in Yunnan, China9 in Lower Pliocene
lignite beds. As lignite, a brown coal intermediate between peat
and bituminous coal, is an excellent preservative, we may hope for
whole skulls and postcranial skeletons from this area. From illus-
trations, the Keiyuan teeth look nearly as hominid as they look
pongid. Professor Woo, who found them, says of the lower first
premolar of his new primate that its outer surface is “worn, as in
other anthropoids, by the posterior inner face of the upper ca-
nine.” Nevertheless, this tooth is short and broad for a pongid.
Whether the type of dental articulation ascribed to it by Woo
could be lost in the evolution of a hominid from a pongid is an
unanswered question. In any case the Dryopithecus teeth from
China seem to be more nearly hominid than those from Europe.
Ramapitliecus, a Possible Ancestor of the Hominids 1
I n 1935 G. E. Lewis and his associates found the jaws and
teeth of many Dryopithecines in the rich fossil-bearing deposits
of the Siwalik Hills in northwest India. They sorted these into
five genera, four of which they named after Indian gods: Siva-
pithecus, Sugrivapithecus, B ramapitliecus, and Ramapithecus.
The fifth was called Paleosimia. Although Ramapithecus has been
7 Le Gros Clark, in i960, called it only “probably” a part of this animal. Clark:
The Antecedents of Man (Chicago: Quadrangle Books; i960), p. 214.
8 Simpson: op. cit., p. 67.
9Ju-Kang Woo: “Dryopithecus Teeth from Keiyuan, Yunnan Province,” VP,
Vol. I, No. 1 (1957), pp. 25-32. Also “New Materials of Dryopithecus from
Keiyuan,” VP, Vol. 2, No. 1 (1958), pp. 38-42.
1W. K. Gregory, M. Heilman, and G. E. Lewis: “Fossil Anthropoids of the
Yale-Cambridge India Expedition of 1935,” CPWP, No. 495 (1938).
Elwyn Simons: “The Phyletic Position of Ramapithecus,” PYPM, No. 57
(1961).
204 The Fossil Record from Lemurs to Swamp Apes
called Upper Pliocene or even Early Pleistocene, it is now as-
signed to the Lower Pliocene along with the other four genera,
and all are roughly contemporary with the other Dryopithecines
mentioned above.
FlG'. 15 JiamaPithecu^ brevirostis. The type specimen of Ramapithecus brevi-
rostts, a Pliocene ape from the Siwalik Hills of Northern India, whose upper molar
and premolar teeth resemble those of man. In this figure Simons has projected
the shape of the whole palatal arc, which appears rounded as in man. ( Drawing by
Simons, 1961 [PYPM, No. 5 7, Fig. 2] with permission.)
The most hominid-looking of these specimens is the piece of
right maxilla of Ramapithecus brevirostis, which contains the first
two molars, both premolars, and the root of the lateral incisor.
The socket of the canine is also preserved, and the lateral wall of
the socket of the median incisor. Simons, who has recently re-
studied this specimen, has reconstructed the palate, and esti-
The Fort Ternan Primate
205
mated the sizes of the missing teeth from the sizes of their sockets
and the space available to each in the tooth-row.
According to this reconstruction the palate is arched, as in
man; the canine was no larger than the first premolar, and was
thick mesiolabially as in man, instead of spatulate, as in apes; and
the ratio between the sizes of the front teeth (premolars and
canines ) and those of the cheek teeth ( premolars and molars ) is
roughly the same as in man, and not as in the apes, which have
relatively large front teeth. Enough of the maxilla is preserved
to show that the upper jaw was more manlike than apelike in its
depth and degree of prognathism. In view of these findings,
Simons has committed himself to the opinion that Ramapithecus
brevirostis was, in fact, a hominid, the first known of his subfamily.
What others will say about this identification remains to be seen.
In any case, India and China may well have been the breeding
places of the Hominidae, either through Ramapithecus brevirostis
or some other species, just as Africa was the cradle of the chim-
panzee and gorilla. The origin of the orangutan is still a mystery,
as no bona-fide orang is known before the Pleistocene.
The Fort Ternan Primate 2
Late in 1961 Leakey turned the putative home of the hominids
back to Africa by discovering a Pontian ( Early Pliocene ) primate
specimen in the orange grove of a white farmer, Fred Wicker, at
Fort Ternan on the Gulf of Kavirondo in Kenya. This discovery
was announced on March 22, 1962. The Argon-40 date deter-
mined at Berkeley is 14 million years, within the accepted span of
the Pontian, and the fauna belongs to that period.
The specimen, like Ramapithecus, consists of a piece of the
right maxilla. It contains the second and first upper molars, the
second upper premolar, and the freshly broken stub of the first
upper premolar. The right upper canine, found separately, has
been glued into the distal cup of its socket.
The specimen is between 4 and 4.5 cm. long, and all the teeth
2 Louis Leakey has asked me to refer to his new find (which I have seen and
handled ) by this name until the official binominal shall have been published. Other-
wise the binominal would be a nomen nudum, and worthless.
206 The Fossil Record from Lemurs to Swamp Apes
are within the human size range. Both molars have the Dryopi-
thecus Y-5 cusp pattern, and show no special features such as a
cingulum, enamel extensions, or enamel pearls ( see p. 358 ) which
characterize certain human races. The size progression from the
molars to the second premolar to the first premolar to the canine
is the same as that found in Homo but not in Australopithecus —
the first premolar cannot have been much larger than the canine.
The canine is short and does not extend downward below the
occlusal level of the other teeth.
The only morphological pecularity of the teeth which I could
observe was a considerable surface relief on the inner or lingual
side of the canine. Like most but not all human maxillae, and no
others, that of the Fort Ternan primate had a canine fossa.
It is easy to speculate on the relationship of this specimen to
Proconsul, Ramapithecus, the living apes, Australopithecus, and
man, but until more details are available, to do so is not only im-
polite but also probably unprofitable.
The Pleistocene Apes of China
As china is the gateway between the Oriental region and
the Palearctic, we should not be surprised to find that during the
Pleistocene a number of higher primate genera in addition to
Homo had established themselves in that country. Among them
is the orang and an animal with huge molar teeth known as
Gigantopithecus blacki,s first described on the basis of molar teeth
3 D. A. Hooijer: “The Geological Age of Pithecanthropus , Megantliropus, and
Gigantopithecus,” AJPA, Vol. 9, No. 3 (1951), pp. 265-81.
G. H. R. von Koenigswald: “Gigantopithecus hlacki von Koenigswald, a Giant
Fossil Hominid from the Pleistocene of South China,” APAM, Vol. 43, Part 4
(1952), PP- 295-325.
W. C. Pei: “Giant Ape’s Jawbone Discovered in China,” AA, Vol. 59, No. 5
(1957), PP- 834-8.
Pei and Y. H. Li: “Discovery of a Third Mandible of Gigantopithecus in Lu-
Cheng (Kwangsi, South China),” VP, Vol. 2, No. 4 ( 1958), pp. 190-200.
W. L. Straus, Jr.: “Jaw of Gigantopithecus,” Science, Vol. 125, No. 3250
(1957), p- 658.
S. M. Gam and A. B. Lewis: “Tooth-Size, Body-Size, and ‘Giant’ Fossil Man,”
AA, Vol. 60, No. 5 (1958), pp. 874-80.
“More Gigantopithecus,” in News and Activities, VP, Vol. 2, No. 1 (1958),
p. 67.
207
Possible Survivals of Chinese Apes
recovered from Chinese pharmacies, where they are sold as tooth-
ache medicine. Between 1956 and 1958 three lower jaws of this
species were removed from a cave in a high cliff in Kwangsi
province. According to Pei and Li, this animal lived in the Lower
Pleistocene, or Villafranchian, earlier than Sinanthropus. How-
ever, the time gap between those two Chinese primates is too short
for Gigantopithecus to have been an ancestor of man, as some
have claimed, whatever anatomical arguments may be produced
to favor such a descent, for in some ways his teeth resembled
man’s more than those of the living apes do. The tooth pattern
was essentially pongid, but the sides of the jaw are convergent
like those of prosimians, and the teeth are worn down all along the
line, indicating a rotary grinding motion, as in hominids. Once
again we are impressed with the capacity of the primates for
parallelism. According to Remane (i960), Gigantopithecus was
definitely a pongid, at the end of a special line. It was less human-
like than a female chimpanzee and about the size of a large
gorilla.
Possible Survivals of Chinese Apes
The Pleistocene ended— if it ended at all — only ten
thousand years ago, a mere yesterday zoologically. It would be
noteworthy if all of the apes of China, the number of genera being
still undetermined, could be shown to have become extinct at the
close of that period. But there is evidence that they did not do so.
For example, the philosopher Hsiin-Tzu, who lived a hundred
years after Confucius, or about 400 b.c., definitely states that an
ape the size of a man and covered with hair lived in the Yellow
River Valley in his day, and also that it stood erect. Furthermore,
the Liang Annals, written in the time of the Warring States,
200 b.c. to a.d. 200, places apes in Sin-Kiang province, north of
Tibet, near the country where the giant panda was first found as
recently as 1930.
A third book, entitled Anatomical Dictionary for Recognizing
G. Heberer: "The Descent of Man and the Present Fossil Record,” CSHS, Vol.
24 (1959), pp- 235-44.
A. Remane: “Die Stellung von Gigantopithecus AAnz, Vol. 24, No. 2-3
(i960), pp. 146-59.
208 The Fossil Record from Lemurs to Swamp Apes
Various Diseases, which originated in Tibet and was published in
Peking at the end of the eighteenth century 4 though it was prob-
ably written earlier, contains a systematic description of the fauna
of Tibet and neighboring regions. Many species of mammals,
birds, reptiles, fish, and so on, are included, and each is illustrated
with a recognizable woodcut. Not one of the animals is fantastic,
composite, or mythical. Among them, in a group of monkeys, a
tail-less, bipedal primate is shown standing on a rock, with one
arm stretched upward. Trilingual captions in Tibetan, Mongolian,
and Chinese designate it as a man-animal. A different and more
detailed illustration appears in an edition of the same book
printed a century later, in Ulan-Bator. In this edition the text
reads: “The wild man lives in the mountains, his origin [this word
probably means habitat] is close to that of the bear, his body
resembles that of man, and he has enormous strength. His meat
may be eaten to treat mental diseases and his gall cures jaundice.”
How, if at all, this wild man is related to the so-called Yeti or
Abominable Snowman remains to be determined, along with its
relationship to the Pleistocene fossil apes of China. If there really
is, or has recently been, a large bipedal primate in central Asia, its
discovery, dead or alive, would be of enormous importance, not
only for primate taxonomy but for its bearing on the theoretical
relationship between the erect posture, tool-making, speech, and
culture.
Hominoids and Hominids 6
From Propliopithecus on (page 196), we have been describing
Hominoids — first hylobatids, or gibbons, then pongids, or great
apes— as distinguished from hominids, or man and kin of men.
Two fundamental features distinguish hominids from their closest
kin, the pongids: posture and teeth. Hominids, by definition,
stand erect and walk with their hands free from the ground. Pon-
4 E. Vlcek : “Old Literary Evidence for the Existence of the ‘Snow Man’ in Tibet
and Mongolia,” MAN, Vol. 59, Article No. 203 (1959), pp. 132-4.
5 The most detailed and authoritative work on this subject, including both
Oreopithecus and Australopithecus, is Heberer: “Die Fossilgeschichte der Homi-
noidea,” in H. Hofer, A. Schulz, and D. Starck: Primatologia, Vol. I (Basel,
1956), pp. 379-560.
Oreopithecus bambolii, the Swamp Ape 209
gids brachiate, walk on their knuckles, or both. Hominids have
small canine teeth that do not project above the line of occulsion
of the other teeth; they have no gap between the upper canines
and first premolars — such a gap is known as a diastema — and the
two lower premolars are more or less the same in shape and func-
tion. Pongids have large, long canines, usually a diastema, and
the first premolar is laterally compressed and has a shearing buc-
cal edge for scissors contact with the upper canine. In each side
of the lower jaw of pongids a hole known as the mental foramen is
located near the lower border of the bone, to clear the long root
of the canine. In hominids this foramen is located higher up,
because the root of the canine is shorter. As we examine the fossil
record in search of hominids, these points must be borne in mind.
Oreopithecus bambolii,6 the Swamp Ape
A recent, much publicized hominid possibility is an animal
found in great abundance in the so-called Pontian fossil beds of
central Italy, which consist, like those of China, of layers of lig-
nite. These beds are attributed variously to the Upper Miocene
and the Lower Pliocene, and cover the period between about 10
and 16 million years ago. This is also the time of the Fort Ternan
primate.
6 Hiirzeler: “Zur Systematischen Stellung von Oreopithecus ,” VNGB, Vol. 65,
No. 1 (1954), pp. 88-95.
Hiirzeler: Oreopithecus hambolii Gervais, A Preliminary Report,” pp. 1—48.
Hiirzeler: “The Significance of Oreopithecus in the Genealogy of Man,” Tri-
angle, Vol. 1, No. 5 (i960), pp. 164-74.
Straus: “ Oreopithecus bambolii,” Science, Vol. 126, No. 3269 (1957) pp
345-6.
Straus: A New Oreopithecus Skeleton,” Science, Vol. 128, No. 3323 (1958)
P- 523-
Straus: “Is Oreopithecus bambolii a Primitive Hominid?” AR, Vol. 132, No. 3
(1958), pp. 511-12.
Straus: “Cranial Capacity of Oreopithecus bambolii,” Science, Vol. 132, No.
3428 (i960), pp. 670-2.
Simons: “Apidium and Oreopithecus,” Nature, Vol. 186, No. 4727 (i960), pp.
824—6.
A. H. Schultz: Einige Beobachtungen und Masse am Skelett von Oreopithe-
cus, ZfMuA, Vol. 50, No. 2 (i960), pp. 136—49.
P. M. Butler and J. R. E. Miles: A Contribution to the Odontology of Oreo-
pithecus” BBMN, Vol. 4, No. 1 ( 1959), pp. 1-26.
E. Bone: “ Oreopithecus bambolii , A Propos du Jalonnement Tertiare de
FHomme,” Q-S., April 20, 1959, pp. 215-46.
210
The Fossil Record from Lemurs to Swamp Apes
First found in the 1860’s and called Oreopithecus, these re-
mains were assigned to various taxonomic categories until Hiirze-
ler reopened the question with new specimens in 1956. Early in
1958 he found a nearly complete skeleton, which at the time of
writing has not been fully described. This animal had been mired
in a forested swamp and covered while still whole, before
Fig. 16 The Skull of Oreopithecus: Hurzeler’s
Reconstruction. ( Drawings after Hiirzeler, i960. )
Fig. 17 The Skull of Oreopithecus:
Drawn from a Photograph. This
picture, drawn from a photograph of
the skull as it lay in its matrix, differs
from Hurzeler’s reconstruction in two
respects: the nuchal crest is higher,
and the mandible is blown out in the
gonial region, as among the leaf-
eating langurs, and suggesting a
specialized diet of soft vegetable mat-
ter. (Drawing after a photograph by
Hiirzeler, i960.)
predators had had a chance to find the body. He was not a moun-
tain ape, as his name implies, but a swamp ape.
The creature apparently stood some 120 cm. or 4 feet high,
about the height of a siamang. Its skull is small, with a length of
125 mm., a breadth of 85 mm., and a capacity of between
275 cc. and 530 cc., which places it in the same brain-body-size
Oreopithecus bambolii, the Swamp Ape 211
ratio as living apes. Although there is no sagittal crest, the supra-
orbital ridges are very heavy. Unlike the faces of apes and early
men, its face is short. However, the zygomatic arch orginates in
the malar bone forward of its position in apes but comparable to
its position in man. There is a suggestion of a nasal spine and the
nasal bones project beyond the surrounding level of the face; both
are manlike features. The symphysis, or sagittal midline, of the
lower jaw is steep but chinless, and the mental foramen highly
placed.
Fig. 18 The Specialized Dentition of Oreopithecus. The tooth on the left, the
upper right third molar, has a small cusp, or conulid, the center of the crown, in
addition to the five cusps characteristic of the Dryopithecines, modern apes, and
hominids. All of its cusps are high and pointed. The upper right second premolar
has five cusps, like a molar, and the two principal cusps are high and pointed.
(Drawings after Butler and Miles, 1959.)
The teeth, which have been thoroughly studied by Butler and
Miles, are similar to man’s in some respects and very different in
others. The canines are small and short, and in eleven of twelve
known jaws a diastema is lacking. Actually, a diastema appears
now and then in human jaws and is sometimes absent in apes.
However, the canines occlude differently from those of hominids
or apes. Both lower premolars are bicuspid and of the same shape;
the shearing edge of the first premolar found in apes is lacking, as
one would expect because of the short canines. The molars are
212
The Fossil Record from Lemurs to Swamp Apes
long and narrow. They have a high cone relief, with a very un-
usual central cone, and thin enamel. The enamel in human teeth
is nearly twice as thick, enabling men to chew more and to live
longer under primitive conditions — unless Oreopithecus ate soft
food.
Butler and Miles find twelve features in which the Oreopithecus
molars differ from those of men, and in most of these twelve our
molars resemble those of the apes. And the Oreopithecus molars
bear no relationship to those of Old World monkeys. The authors
conclude: “This peculiar combination of primitive and special-
ized characters seems to indicate that Oreopithecus is the termi-
nal form of an independent phyletic line that extended back prob-
ably into the Oligocene.” 7
While Hiirzeler has concentrated on the preparation of the
skeleton of Oreopithecus and Bulter and Miles have studied its
teeth, Schultz has concerned himself with the postcranial skele-
ton.8 It was the appearance of the pelvis and limb bones which
initially led Hiirzeler and de Terra to the widely publicized
theory that the animal might have already assumed the erect
posture. Schultz has effectively undermined this concept. He
took twenty measurements of the pelvic and limb bones, exclusive
of the hands and feet, and calculated fourteen indices. Then he
compared this data with similar measurements and indices from
ten Old World monkeys of more or less the same size, a gibbon,
a siamang, a male and a female orang, two male chimpanzees, a
female mountain gorilla and a male lowland gorilla, a Negro, a
Hawaiian, and a European.
The animal had a trunk height ( supersternale to symphysion)
of about 460 mm., which is in the range of the largest Old World
monkeys, the orang, and the chimpanzee. Judging by its build it
weighed about 40 kilograms (88 pounds), a weight equaled
among the Old World monkeys only by the largest baboons, and
within the orang-chimpanzee range. Its humerus was as long as
those of chimpanzees and men, and longer than those of all
Old World monkeys. Its femur was shorter than those of Old
World monkeys and all apes but the orang. The head of the hu-
7 Butler and Miles: op. cit.
8 Schultz: “Einige Beobachtungen. . . .”
Oreopithecus bambolii, the Swamp Ape 213
merus is much wider than that of the femur, which indicates that
the body was supported by the arms more than by the legs; and
the humerus is 122 per cent as long as the femur, a proportion
found in the siamang, orang, and lowland gorilla. Neither the
bones of the forearm nor those of the lower leg were particularly
Fit;. 19 The Pelvis and Femora of Oreopithecus. The top drawing represents
the pertinent bones of a langur, the middle one of Oreopithecus, and the
bottom one of a chimpanzee. Note the relative shortness and breadth of the
Oreopithecus pelvis, which in this respect appears hominid, and the shortness and
stoutness of the femora, which in this sense are pongid. (Drawings after Schultz
i960. )
elongated. Although a brachiator, Oreopithecus was not ex-
tremely specialized in the proportions of its limb segments. That
it was arboreal is to be expected because most of the fauna of
flooded forests is either tree-borne or aquatic.
Several bones indicate that the animal had a stout body. The
femur is thick in proportion to its length, the lumbar vertebrae are
heavy, and the ribs are large. The shape of the ribs in particular
is apelike rather than monkeylike, in that the dorsal arm of the rib
214 The Fossil Record from Lemurs to Swamp Apes
lies at an obtuse angle to the main body of the bone, whereas in
monkeys the angle is acute. Five lumbar vertebrae were present,
compared to between six and eight in monkeys, between four and
six in gibbons and men, and between three and five in the great
apes.
The form of the pelvic bone ( os coxae ) is of particular interest
as it led to the early conception that Oreopithecus stood erect.
The pelvis as a whole is broad, but no broader than in apes. How-
ever, the ilium of the ape’s pelvis is long and high; that of Oreo-
pithecus is shorter, as in man and in the larger Old World mon-
keys. The pubic symphysis is short and straight, as among apes
and men, but not monkeys; the ischium small and short, as among
apes and men, and not long as in monkeys, gibbons, and one
extinct hominid that will be discussed in the next chapter ( Aus-
tralopithecus). Apparently Oreopithecus had not yet developed
the pelvic peculiarities of the other large brachiators.
Schultz concludes that Oreopithecus is a catarrhine, but that it
does not belong to the Cercopithecidae because of peculiarities
of both teeth and body. It belongs to the superfamily of Homi-
noidae, but not to the gibbons ( Hylobatidae ) because of its
teeth and ischium, its short femur, and short lower arm bones. As
for the family of Pongidae, its kinship is near but not exact: five
lumbar vertebrae is a high number for this group, its ilium has not
become elongated, and its teeth are aberrant. It definitely does
not belong with the Hominidae; there is no evidence of upright
posture superior to that of the larger apes. In its limb bones it is
closest to the gorilla, which is the least efficient brachiator among
living apes.
In 1916 Schwalbe called the Oreopithecidae an extinct family,9
and it is either that or a connecting form between the Pongidae
and Hominidae. This will be clearer when more evidence is
available. Kalin in 1955, 1 Thenius in 1958, 2 and Butler and Miles
in x959 3 have all confirmed Schwalbe’s theory. One may conclude
9 G. Schwalbe: “tlber den fossilen Affen, Oreopithecus hamholii,” ZfMuA, Vol.
19 (1916), pp. 149-254.
1 J. Kalin: Zur Systematik und evolutiven Deutung der hoheren Primaten,”
Experientia, Vol. 11 (1955), pp. 1-17.
2 E. Thenius: “Tertiarstratigraphie und tertiare Hominoidenfunde,” AA, Vol.
22 (1958), pp. 66-77.
3 Butler and Miles: op. cit.
Fossil Primates and Human Evolution 215
with Schultz that Oreopithecus adds to the number of known
forms of Hominoidea, and constitutes further evidence “of the
extraordinary variability and plasticity of this group, to which
man belongs.” 4
Fossil Primates and Human Evolution
At the beginning of this chapter we proposed to test, on the
scale of time, the conclusions of comparative anatomy, physi-
ology, cytology, and parasitology as to the degrees of kinship be-
tween man and his fellow primates. According to these conclu-
sions, man fell closest to the living apes; but these findings are still
unconfirmed. To date we have no certain ancestor earlier than the
Pleistocene, although Ramapithecus brevirostis and the Fort
Teman primate are distinct possibilities, and Oreopithecus
bambolii has warm champions.
We also hoped to ferret out some evolutionary rules that might
cast a few rays of light on man’s evolution into geographical races.
Zoologically this is relatively minor, but to us it is important. De-
spite the meager paleontological representation of primates and
despite many wide gaps in the record, we discovered a few en-
lightening continuities.
The primates appeared early in the history of mammals as a
very generalized order of tiny animals, arboreal, virtually omnivo-
rous but with an accent on animal proteins, and reproductively
primitive. As prosimians they showed a remarkable adaptive ver-
satility. Some became nocturnal, some acquired a slowed-down
metabolism, some lost their tails, some poked the spines of their
dorsal vertebrae through their skins as weapons, and some grew
chiseling incisors like those of rodents. From these adaptations
several superfamilies passed into a higher adaptive grade, that of
monkeys (the simian), and they made this transition independ-
ently in both hemispheres. Both groups acquired stereoscopic
color vision and rapid locomotion in the trees, through brachia-
tion.
It is possible that in South America the marmosets and the
cebus monkeys crossed the frontier from the prosimian to the
4 Schultz: op. cit., p. 148; translation mine.
2l6
The Fossil Record from Lemurs to Swamp Apes
simian grade independently also, just as four different kinds of
reptiles once became mammals. It is equally possible that the Old
World monkeys and the ancestors of the apes and men made the
same transition separately, although this too has not been proved.
Parallel evolution, in separate but genetically similar populations,
is a primate commonplace, and a zoological commonplace as well.
Another fact of outstanding significance evidenced in the pale-
ontological record is that in the early epochs of the Cenozoic, tens
of millions of years ago, all the primates were small, ranging in
size from mice to squirrels and cats. In the Miocene only Procon-
sul, as far as we know, was bigger than a gibbon. In the evolution
of the different primate lines the principle of allometry, or the
shift of bodily proportions with growth, has been at work. In the
elephants the leg and foot had to become columnar to support
the animal’s increasing weight. In a brachiating primate the hand
needed tendinous supports once a certain weight threshold had
been reached. In a bipedal primate the pelvis, legs, and feet
needed special modifications for a large animal that a small ani-
mal did not need.
Several families of primates learned to brachiate independently,
and it is possible by the same token — although we cannot prove
this — that more than one subfamily or genus independently be-
came bipedally erect. In different families, some genera have
grown more intelligent than others. In fact, although a definitely
identified early ancestor of man still eludes us, we have dis-
covered a pattern that we might look for when we turn to the
history of our own egocentric species, as well as that of other prob-
ably less articulate, and extinct, species and genera in our own
family. Because changes of all magnitudes, which eventually
designate species, genera, and higher taxonomic categories, be-
gin at a subspecific level, what we have learned in this chapter
provides a background for a realistic evaluation of the evolu-
tionary parallelism that exists and has existed between various
geographical races of man at successive stages of human evolu-
tion.
7
THE EARLIEST HOMINIDS
W
The Origin of the Hominids
e have traced in some detail the family histories of
all the primates, except the Hominidae, from the Paleozoic to the
present. That of the Hominidae has been postponed until now for
two reasons. They include all fossil and living men, whose evolu-
tion into races constitutes the main subject of this book, as well as
a subfamily of related manlike creatures, the Australopithecines.
They are the only primate family lacking a known, proven ances-
tor who lived before the Pleistocene. Not a trace of the Hominidae
has yet been found in a deposit incontestably older than the end
of the first half of the Lower Pleistocene.1 Yet between that time
and the beginning of the Middle Pleistocene their bones or tools
or both were deposited in several sites scattered all the way from
Algeria to South Africa and Java. What happened to the Homini-
dae during this earliest known period of dispersal is pertinent to
the study of human racial origins because the differentiation of
races may have begun by or during that time.
Did the Hominidae exist before this dispersion took place, and
if so, where? These questions cannot be answered conclusively on
the basis of existing information: entire families and subfamilies
of primates remained hidden over vast geological periods. Gaps of
over 30 million years separate the tree shrews and tarsiers from
their last known ancestors, and the chimpanzee and gorilla are
parted from Proconsul , an ape whose name will appear frequently
in this chapter, by 25 million years.
1 Except possibly for one Australopithecine fragment from Tchad, to be de-
scribed later, and unless Ramapithecus and the Fort Teman primate are hominids.
2l8
The Earliest Hominids
Of 118 families of mammals living today,2 ig first appeared dur-
ing the Eocene or earlier, 22 during the Miocene, 13 during the
Pliocene, 12 during the Pleistocene, and 27 in the Recent epoch.
During the Pliocene, which lies in the middle of this progression,
the several families of large land mammals evolved, including
elephants, rhinoceroses, and hippopotamuses.3 In contrast, most
of the land mammals that first appeared as families during the
Pleistocene or Recent epoch are small creatures: rodents, insec-
tivores, and prosimians. The Pliocene, therefore, was a reasonable
peiiod for a family of fairly large-bodied animals with only one
living genus, the Hominidae, to have evolved in. If our family
came into being during the Upper Pliocene or during the initial
phase of the Lower Pleistocene, no paleontological precedent or
protocol was violated.
The bulk of the anatomical and physiological evidence re-
viewed in the last two chapters strongly suggests that our ances-
tors evolved from the same primate stock as the chimpanzees,
gorillas, and orangs. We differ from these three apes and from
the Early Miocene ape, Proconsul, in three principal respects:
locomotion, brain size, and dentition. Neither locomotion nor
brain size is significant as far as our relationship to the living
pongids is concerned. The apes might have begun to brachiate
before our ancestors quit their company, or our ancestors might
have begun to brachiate, with the apes, before walking erect.
And the brains of all evolutionary lines of primates must have
been small at the beginning. But dentition is a more serious barrier
to kinship with the living pongids, all of which have interlocking
canines and shearing lower first premolars. The shift from the
shearing and crushing type of teeth peculiar to all living and most
fossil apes to our grinding type was a dramatic one, which may or
may not have taken place more than once.
Pending the final report on the Fort Ternan primate, the lead-
ing candidates for the title of Pliocene ancestor of the hominids
will still be the Dryopithecinae found in the Siwalik deposits of
2 Calculated from G. G. Simpson’s list of 1945.
3 The hippopotamus is a land animal in the sense that it comes out of the
water at night to feed on dry land.
2ig
The Origin of the Hominids
northern India in the 1930’s by G. E. Lewis and his associates.4
Because these consist of teeth and jaws alone we do not know
whether these apes brachiated or walked, nor how large their
brains were. We know only that for some reason their jaws were
shorter than those of other pongids and the teeth of some of them
were no larger than those of the living genus Homo and were, to a
certain extent, manlike in form. Others had larger teeth. The
body sizes of all these genera are unknown.
In the genus Ramapithecus an evolutionary sequence, pointing
in the hominid direction, may be traced from R. hariensis of the
Lower Pliocene to R. brevirostis of the Upper Pliocene and initial
Pleistocene. In this sequence the molars grow wider (labiolingu-
ally) than they are long (anteroposteriorly). The premolars be-
come bicuspid, and although the lower first premolar is still sec-
torial, the canines are small, and there is no diastema, or gap
between the upper canines and the upper lateral incisors. The
incisors rise steeply from both jaws, and the alveolar prognathism
is less than in some living human beings.
Proconsul himself, who sired the gorilla and chimpanzee, was
an African Dryopithecine, and the earliest member of his family
yet known. He was related, either as an ancestor or as a cousin,5 to
the Dryopithecines of Europe and Asia, which lived from mid-
Miocene to Early Pleistocene and which included the god-apes of
India, Gigantopithecus, and the as yet unidentified ancestor of
the orang. An ape identified in 1951 by LeGros Clark and Leakey
by its teeth as Sivapithecus africanus, attributed to the Lower
Miocene of Kenya, may have been the link between the African
and Eurasiatic Dryopithecines. Therefore Proconsul could have
4 G. E. Lewis: “Preliminary Notice of the New Man-like Apes from India,”
AJSc, Ser. 5, No. 27 (1934), pp. 161-79.
W. K. Gregory, M. Heilman, and G. E. Lewis: “Fossil Anthropoids of the Yale
Cambridge India Expedition of 1935,” CIWP, No. 495 (1938).
See also G. Heberer: “Die Fossilgeschichte der Hominoidea,” Primatologia
(Basel, 1956), pp. 379-560.
5 In the sense of G. G. Simpson, who wrote, in 1945: “The -}- Dryopithecinae
are probably a very heterogeneous group which represents a stage in primate evo-
lution rather than a single phylum and its branches. Thus the different Ponginae
probably arose from different Dryopithecinae so that the separation of the sub-
families is not phylogenetic classification, but the true phyla are not really dis-
tinguished at present.” (p. 188)
220
The Earliest Hominids
been our ancestor, either through an African line including the
Fort Ternan primate and distinct from that of the gorilla and
chimpanzee, or through an Asiatic line that left Africa in the
Miocene or Pliocene and returned in the Lower Pleistocene. This
geographical problem cannot be decided on present evidence.
Australopithecus and Homo
W ii e r e v e r it originates, a new family can arise when a group
of animals adopts a new ecological position by a radical change
of behavior, as, in the case of man, by walking erect, using tools,
talking, and seeking food on the ground in groups. But it takes
time for a new family to branch into a number of genera and for
genera to give birth to species, either by succession, branching, or
both; and our family, as we know it, has had very little time.
The known Hominidae are divided into two genera, Aus-
tralopithecus and Homo. Australopithecus lived during the
Lower Pleistocene, with a slight overlap into the Middle Pleisto-
cene. Except in Java, Homo is so far definitely known only from
the beginning of the Middle Pleistocene onward. The question
thus arises, is Homo descended from Australopithecus by evolu-
tion through succession, i.e., by phyletic evolution, or did the two
genera arise from a common ancestor through branching, after
which our genus replaced its brother?
Both theories have warm champions, and the question is not
likely to be decided immediately. So that we may understand the
problem as clearly as possible I shall devote the rest of this chap-
ter to the first known phase of hominid history, the Lower Pleisto-
cene, and particularly to the Australopithecines.6
6 The name Australopithecinae, denoting a subfamily, was coined by Gregory
and Heilman in 1939. Whether these animals actually form a subfamily or just a
genus is a matter of opinion. Gregory and Heilman: “The Dentition of the Ex-
tinct South- African Man-Ape Australopithecus ( Plesianthropus ) transvaalensis
Broom, A Comparative and Phylogenetic Study,” ATM, Vol. 29 (1939), pp.
339-73- To match the term Australopithecine, the word Hominine is sometimes
used for man.
The Lower Pleistocene
221
The Lower Pleistocene
The Lower Pleistocene is the name given the first half of the
Pleistocene epoch; it is believed to have begun about one million
years ago and to have ended about one half million years ago.
Lower Pleistocene deposits cannot be easily distinguished from
the underlying Pliocene beds everywhere that both occur, but in
some places an abrupt soil change caused by uplift and erosion
marks the Plio-Pleistocene threshold and in others no Pliocene
underlies the Pleistocene strata at all.
With the advent of the Pleistocene the climate cooled re-
peatedly in certain parts of the earth, and in others there were
alternating periods of heavy rainfall and drought. New mountains
rose and old ones increased their stature, volcanoes spouted lava
and dust, and sea levels rose and fell, as the earth’s crust buckled
and waters of the oceans were first imprisoned in icecaps and then
released, three such cycles occurring in a row. But the icecaps
were a special feature of the second half of the Pleistocene. Dur-
ing the Lower Pleistocene mountain glaciers formed in stream
beds several times in different places, but no ice accumulated on
continental land masses. When the first icecaps appeared on
continental land masses, the Middle Pleistocene had begun.7
In certain critical places the point in time at which the Pleisto-
cene began is defined by the appearance of cold-adapted molluscs
in previously warm waters.8 The change in molluscan geography
is matched, on land, by the appearance of modern genera and
subfamilies of horses, cattle, elephants (in the form of mam-
moths), and camels. These animals spread rapidly and widely
in the Palearctic, Nearctic, Oriental, and African regions.
The fauna to which they belonged is called Villafranchian, after
a site in Italy. In the original sense the name meant a particular
assemblage of mammals, but in various places these animals con-
tinued to evolve, so that many old species were replaced by new
7 See J. K. Charlesworth: The Quaternary Era (London: E. Arnold & Co.;
1957)-
8 See F. C. Howell: “The Villafranchian and Human Origins,” Science, Vol.
130, No. 3379 (1959), PP- 831-44.
222
The Earliest Hominids
ones. Some genera even replaced others. The Lower Pleistocene,
with its challenging alternations of climate, was a time of rapid
mammalian evolution. Many authors use the term Villafranchian
as a synonym for Lower Pleistocene. This practice has caused
some confusion because many local faunas of the middle and
latter parts of the Lower Pleistocene are Villafranchian only in a
general and derivative sense.
Lower Pleistocene sequences of the Old World, some geological,
some faunal, and some both, are best known from Europe, Pales-
tine, India, Java, China, and several parts of Africa. In western
Europe a succession of mountain glaciers and associated drops in
temperature produced first a cold phase, then a so-called Tiglian
cool interglacial, then the two mountain glaciers known as Giinz I
and Giinz II, and finally the Cromerian Interglacial, which was
followed by the beginning of the Middle Pleistocene. In Central
Europe three local mountain glaciations, called Donau I, II, and
III, occupied the same general time span.9
In Palestine two successive earth movements, or riftings,
opened the crack that created the Jordan Valley and the Dead
Sea. The twisted strata and dislocated blocks so formed mark the
thresholds between the Pliocene and Lower Pleistocene and be-
tween the Lower and Middle Pleistocene.
In India the Pliocene deposits of the Siwalik Hills, so rich in
pongid fossils, are overlaid by two successive Lower Pleistocene
levels, first the Tatrot, then the Pinjor. The Tatrot is Early Lower
Pleistocene, and the Pinjor is contemporaneous with the first of a
series of four Himalayan glaciations, corresponding to Giinz in the
Alpine series. Although much alike, the Tatrot and Pinjor faunas
9 As I write, this classification is changing. B. Kurten has proposed a new way of
dividing the first half of the Pleistocene which makes excellent sense. In it the
Lower Pleistocene is synonymous with the Villafranchian, which F. Clark Howell
calls the Basal Pleistocene. Thus the Giinz glaciations, the Cromerian Interglacial,
and the Mindel-Elster glaciation (including the Cortonian Interstadial between
Mindel I and Mindel II) together become the Lower Middle Pleistocene. Then the
Second, Great, or Holstein Interglacial and the following Riss, Saale, or Third Gla-
ciation become the Upper Middle Pleistocene. What is left remains Upper Pleisto-
cene, as before. Although this scheme has much merit, it needs to be generally
accepted (as it probably will be) before I can use it in a book. B. Kurten: “The
relative ages of the Australopithecines of Transvaal and the Pithecanthropines of
Java, in G. Kurth: Evolution und Hominisation (Stuttgart: Gustav Fischer Verlag;
1962), pp. 74-80.
The Lower Pleistocene 223
are differentiated by the presence or absence of certain species.
For example, Ramapithecus, the Dryopithecine ape with the most
humanlike dentition so far described, lived on into the Tatrot but
is not found in the Pinjor.
Most of Java was submerged during the Pliocene, but during
the Pleistocene the land rose, starting at the western end. Geologi-
cal deposits in the western part of the island are clearly Lower
Pleistocene, like those of the Siwaliks. In the eastern part of the
island the earliest fauna is found in the Djetis beds, which have
been called contemporary with either the Cromerian Interglacial
or the beginning of the first Mindel glaciation of the initial Mid-
dle Pleistocene. These beds contain a so-called Sino-Malayan
fauna, including the orang, gibbon, and two hominids, all of
which are believed to have originated in south China and to have
reached Java via the outer ring of islands.
In south China the bones of this fauna are cemented in blocks
of breccia preserved in rock crevices. Because breccia can have
formed at several different times, these fossils have been dated on
the basis of their first appearance in stratified deposits elsewhere,
in this case the well-known Siwalik Hills. There the Sino-Malayan
fauna is absent from the Pinjor beds. For that reason Hooijer and
others, including Howell, call the Djetis fauna Early Middle Pleis-
tocene, whereas von Koenigswald, who did much of the original
research in Java, has consistently stipulated a Late Lower Pleisto-
cene date.1
At present von Koenigswald seems to be ahead in this argu-
ment. In 1961 he obtained two tektites (glassy nodules from outer
1 D. A. Hooijer: “Fossil Mammals and the Plio-Pleistocene Boundary in Java,”
PKAW B, Vol. 55, No. 4 ( 1952), pp. 436-43.
Hooijer: “The Lower Boundary of the Pleistocene in Java and the Age of
Pithecanthropus,” Quaternaria, Vol. 3 (1956), pp. 5-10.
Hooijer: “The Correlation of Fossil Mammalian Faunas and the Plio-Pleisto-
cene Boundary in Java,” PKAW B, Vol. 60, No. 1 (1957), pp. 1-10.
Howell: “The Age of the Australopithecines of Southern Africa,” AJPA, Vol.
13, No. 4 (1955), PP- 635-62.
Howell: “The Villafranchian and Human Origins,” Science, Vol. 130, No. 3379
(1959), PP- 831-44.
Heberer: op. cit., pp. 379-560, 528.
Von Koenigswald: “Remarks on the Correlation of Mammalian Faunas of
Java and India and the Plio-Pleistocene Boundary,” PKAW B, Vol. 59 (1956),
pp. 204-10.
224
The Earliest Hominids
space) from two different deposits in central Java, taken from
beds of the Trinil fauna — the one following the Djetis, with which
we are here concerned. He submitted both samples to the atomic
laboratory of the Max Planck Institute in Heidelberg. There
W. Gentner and H. J. Lippolt tested them by the Argon-40
method, which will be described in tire following chapter. The
two samples gave almost identical results. The Trinil beds were
laid down about 500,000 years ago. That is the conventional date
The Lower Pleistocene
225
of the beginning of the Middle Pleistocene. The Djetis beds, being
older, are therefore of Late Lower Pleistocene date.2
In northern China, where no atom-age dating has yet been
done, sequences have been worked out. The Pliocene was warm
and mainly dry. The Pleistocene, which started with earth move-
ments, was wetter and cooler. The Lower Pleistocene deposits
consist first of basal conglomerates, then of a series of sands,
marls, and clays known as the Lower Sanmenian, and finally of a
bed of sands and silts containing an Asiatic version of the Villa-
franchian fauna. Next comes an erosion surface caused by addi-
tional earth movements, and then the Upper Sanmenian red
loams, which are Middle Pleistocene.
In East Africa the giant earthquakes that cracked out the Jor-
dan also split open the Rift valleys and lowered the lake beds.
Among the flanks of the Rift faces, gullies have been eroded
through 300 feet or more of Pleistocene deposits, some of which
are seated on beds of basalt. There is no question about locat-
ing a Pliocene-Pleistocene border there because no Pliocene
deposits have yet been identified. As they consist of alternate
layers of lake-bottom accumulations and volcanic ash, some of the
Lower Pleistocene beds are very thick. But these materials can ac-
cumulate rapidly; the thickness of the beds is therefore not an
accurate indication of the passage of time.
Five key East African sites have yielded local Lower Pleisto-
cene faunas: Kaiso, Omo, Kanarn, Laetolil, and Olduvai. Of these
Kaiso is considered to be the oldest. Omo and Kanam, roughly
contemporaneous, come next and overlap those below and above
them in the time scale. Laetolil is the next to youngest and Oldu-
vai the most recent.3 At the end of the Lower Pleistocene the East
African climate, which had been moist, grew very dry, and during
the drought the top of the Lower Pleistocene deposit at Olduvai
2 Von Koenigswald, W. Gentner, and H. J. Lippolt: “Age of the Basalt Flow
at Olduvai, East Africa,” Nature, Vol. 192, No. 4804 (1961), pp. 720-1. “Das
absolute Alter des Pithecanthropus Erectus Dubois,” in Kurth: Evolution und
Hominisation (Stuttgart: Gustav Fischer Verlag; 1962), pp. 112-19. Also personal
communication from von Koenigswald, December 20, 1961.
3 In the East African faunas Ewer found the following percentages of living
species: Omo, 21%; Laetolil, 29%; and Olduvai, 36%. The Kaiso fauna had only
thirteen species, too few to be statistically significant. R. F. Ewer: “Faunal
Evidence on the Dating of the Australopithecinae,” PTPA, 1957, pp. 135-42.
226
The Earliest Hominids
Gorge weathered away. We do not know how much of the Oldu-
van faunal deposits were lost at that time.
In North Africa several sites contain Lower Pleistocene fauna,
but of a relatively late date, comparable to those of the East
African locations. The best known is Ain Hanech (more properly
Hanash) or Snake Spring, near St. Arnaud in the Setif plateau,
department of Constantine, Algeria.
In South Africa the picture is confused by the influence of the
antarctic air masses, which make the local sequence partly inde-
pendent of the glacial and pluvial systems farther north. Here the
chronology of the Lower Pleistocene is based largely on fauna,
some of which seems older than the East African series because it
contains a larger number of extinct species. When we study the
Australopithecines, this fact will assume a considerable impor-
tance.
The New Dating for the Lower Pleistocene
Until August, 1961, most geologists, paleontologists, and
anthropologists were content to accept, at least provisionally, the
date of about one million years ago for the beginning of the Villa-
franchian. Then the National Geographic Society dropped a
bombshell in a press release, following it with a magazine article
in October.4
The bombshell was a new date for the Zinjanthropus level in
Bed I of Olduvai Gorge, Tanganyika, determined by the newly
discovered Argon-40 method, the same one used by von Koenigs-
wald’s associates on Javanese samples. J. F. Evernden and G. H.
Curtis of the University of California at Berkeley collected several
samples of volcanic materials at the Gorge and measured them for
Argon-40 content. The dates provided by these samples, taken
from the Zinjanthropus level of Bed I, ranged from 1,570,000 to
1,890,000 years, with an average of 1,750,000 years ago.
The confusion which the publication of these dates caused was
somewhat allayed five months later when von Koenigswald and
Lippolt announced that the basalt underlying Bed I was only
4 G. H. Curtis: “Clock for the Ages: Potassium Argon,” NG, Vol. 120, No. 4
(1961), pp. 590-2.
The Evidence of Tools and Fire 227
1,300,000 years old. However, doubt was revived after a few
weeks, in March 1962, when Leakey declared, on the basis of
new tests at Berkeley, that the basalt was really 4 million years
old.
Lippolt tested two chopping tools made of basalt which von
Koenigswald had collected in Beds I and II, and a piece of the
underlying basalt chipped off by Oakley. All gave a date of
1,300,000 years. Meanwhile Curtis and Evernden, according to
Leakey, got dates of nearly 4 million years from the same under-
lying basalt, but from different samples. As Leakey himself has
suggested, it seems likely that the basalt was laid down more than
once and that different layers have different dates. In this sense,
everyone is right.
Either the soils of Bed I were laid down by wind and water
from older volcanic deposits, or else the different minerals sent to
laboratories from Olduvai Gorge accumulate Argon-40 at differ-
ent rates, as Straus and Hunt have suggested.5 Otherwise the
post- V illafranchian part of the Lower Pleistocene lasted much
longer than had been supposed, not only in Africa but through-
out the world. For the purposes of this book I shall adhere to
the conventional chronology, at least in this edition.
The Evidence of Tools and Fire in the Lower Pleistocene
The Hominidae are distinguished from the other primate
families by a behavioral characteristic that can be determined
archaeologically — the manufacture of stone tools. Whenever
hominid bones have been found in undisturbed habitation sites,
tools are there also. Other tools of equal age have been found in
disturbed and undisturbed sites lacking hominid remains. This
does not mean that all hominids made stone tools, only that there
is no proof to the contrary.
Lower Pleistocene tools whose age can be definitely certified
have so far come only from North and East Africa. The principal
5 Von Koenigswald: “Das absolute Alter des Pithecanthropus Erectus Dubois,”
in Kurth: Evolution und H ominisation, pp. 112-19. Also personal communication
from L. S. B. Leakey, March 24, 1962. W. L. Straus, Jr., and C. B. Hunt: “The
Age of Zinjanthropus,” Science, Vol. 136, No. 3513 ( 1962), pp. 293-5.
228
The Earliest Hominids
site in North Africa is again Ain Hanech,6 situated in or on the
edge of an old lake deposit containing a Late Lower Pleistocene
fauna similar to that found at Olduvai.7
In East Africa tools have been found with a Kaiso fauna at
Kanyatsi, Uganda; at the faunal sites of Omo and Laetolil; and at
Olduvai itself. In South Africa a few tools have turned up in the
more recent of the Australopithecine cave sites. They are probably
no older than those from East Africa, if as old. These African sites,
with the exception of the latest South African one, are older than
the oldest known specimens of Homo in Africa, and the fossils
that have been found with tools probably, if not definitely, belong
to the genus Australopithecus.
The tools that have been found at single sites in large enough
numbers to constitute complete industries follow a definite pat-
tern. Some are simply oval, water-rounded pebbles split crosswise,
lengthwise, or diagonally. Others are single-edged choppers or
double-edged chopping tools, and still others are crude, simple
flakes. In some of the North and East African sites stone balls have
been found. These have fancifully been called bolas stones, but
the most perfect ones are much more plausibly stones especially
shaped for accurate throwing. Anyone who has played baseball
knows that it would be easier to hit an animal with a stone shaped
like a perfect sphere than with a shapeless piece of rock.
Wherever these implements have been dated, by faunal as-
sociations or otherwise, they have come from the later part of the
Lower Pleistocene. As they have been found nowhere at the base
of the Pleistocene, they may be considered a Middle or Late
Lower Pleistocene invention, and as far as we know an African
one.
In Eurasia most if not all of the tool-bearing sites or groups of
sites so far found, from England to the Philippines, cannot be
shown to be older than the Cromerian-Mindel threshold that
marks the end of the Lower Pleistocene, with possible exceptions
in France, India, and Malaya.
In 1959 two French archaeologists, R. Agache and F. Bordier,
6L. Balout : Prehistoire de VAfrique du Nord (Paris: Arts et Metiers Graphi-
ques; 1955), pp. 159-73-
7 C. Arambourg: “L’Hominien Fossile d’Oldoway,” BSPR, Vol. 57, Nos. 3/4
(i960), pp. 223-8.
The Evidence of Tools and Fire 229
while excavating a small trench on the highest terrace of the
Somme River near Moulieres, uncovered a Lower Pleistocene
deposit apparently of Tiglian age. In it they found a tooth of the
Villafranchian horse Equus stenonis, several flint flakes at least
one of which they identified as an implement, and what seemed to
be a hearth.8 Additional work needs be done there before this
discovery can be evaluated.
In Palestine, also in 1959, a bulldozer operator accidentally un-
covered what seemed to be a habitation site in a Lower Pleisto-
cene outcrop at Tell Ubeidiya, Israel, near the southern shore of
Lake Tiberias and just west of the Jordan River. In addition to
very fragmentary hominid remains they found chipped balls,
choppers, chopping tools, and several flakes, constituting the
industry seen at Ain Hanech and Olduvai. Although the exact age
of these finds remains to be determined, they are probably no
older than the two African sites, that is, post-Villafranchian.
In Northern India typologically good Lower Pleistocene im-
plements of the so-called pre-Soan industry have been found with
a fauna of Cromerian age in the gravels of the Second Himalayan
Glaciation. These should be at least as old as the Lower-to-Middle
Pleistocene threshold.1
In Malaya also, tools have recently been found in gravels of
probably the same age, Cromerian or earliest Mindel, which could
make them as old as the Djetis beds of Java.2 These tools are
R. Agache and F. Bordier: Decouverte de Silex Apparemment Tallies a un
Equide Archaeique de Type Villafranchien dans la Haut Terrasse Superieure de
la Somme, CRAS, Vol. 248, No. 3 (1959), pp. 439—40.
9 M. Stekelis, L. Picard, N. Schulman, and G. Haas: “Villafranchian Deposits
near Ubeidiya in the Central Jordan Valley (Preliminary Report),” BRCI, Vol.
9-G, No. 4 (i960), pp. 175-84.
These remains are said to come from a stratum containing Melanopsis in-
vertebrate fauna. The Melanopsis stage of the Lower Pleistocene is a very
early Lower Pleistocene lake-bed formation, entirely under water, which could
hardly have been a living floor at the time it was formed, although hominids could
have camped on the shore. The mammalian fauna has not yet been identified in
enough detail to pinpoint the exact stage of the Pleistocene to which it belongs,
but the preliminary report contains no genus or species name inconsistent with a
Lower Pleistocene date.
1 R- E- M. Wheeler: Early India and Pakistan (New York: Frederick A. Prae-
ger; 1959), PP- 34-62.
Ann Sieveking: The Paleolithic Industry of Kota Tampan, Perak, North-
west Malaya, AP , Vol. 2, No. 2 (i960), pp. 91—102.
230
The Earliest Hominids
mostly single-faced choppers. Siam and the Philippines are begin-
ning to yield similar industries, so far undated. The earliest indus-
tries of the Far East differ in detail from those of the West. Their
principal tool is the single-faced chopper, and there are no
flaked balls.
The currently popular theory that tool-making began in Africa
in the Late Lower Pleistocene and spread to Europe and Asia only
at the beginning of the Middle Pleistocene has not yet been dis-
proved, but it faces many challenges as more and more archae-
ological research is carried out in India, southeast Asia, and Indo-
nesia (in the ethnic, not political, sense). However, it seems likely
that noithern China, whose Lower Pleistocene beds have been
carefully explored, was uninhabited until the Middle Pleistocene.
Tools and fire are the unique possession of man. Except for the
unconfirmed discovery of Agache and Bordier, however, no evi-
dence of fire whatever has been found in any Lower Pleistocene
site, either in the form of charcoal, charred bones, or discolored
stones. And, as we shall see shortly, the teeth and jaw muscles of
most if not all known Lower Pleistocene hominids were big
enough and strong enough to masticate raw food, including meat.
Geography and Numbers of Early Hominids
As one would expect, the sites containing Lower Pleistocene
tools outnumber those containing early hominid bones. An ani-
mal which makes stone implements discards thousands of chips
and tools in his lifetime, and, as stone is inedible, these artifacts
usually stay where he has left them, unless they are moved by
water or ice. He himself has only 180-odd bones in his body and
all of him except his teeth is edible. Unless he happens to drown or
sink in quicksand his remains will most likely be dragged away,
dismembered, and digested. Several of our early hominid skele-
tons, therefore, were found under what once was water.
Ten sites and possibly also a Chinese drugstore 3 have furnished
remains of a maximum of seventy-five hominid individuals, pre-
3 Ralph von Koenigswald found some fossil teeth, which he considers to be
Australopithecine, in the same Chinese pharmacies in which he also found Gi-
gantopithecus teeth. See page 358.
The South African Australopithecines 231
sumably Australopithecines, which will be described in this chap-
ter. These remains have in common the fact that they were
found in Lower Pleistocene deposits, or, if they are younger, that
they cannot be definitely called Homo. Their geographical distri-
bution is as follows.
TABLE 4
THE DISTRIBUTION OF
EARLY HOMINIDS
Region
Site
Number
South Africa
Taung
1
Sterkfontein
21 ca.
Makapansgat
5 ca.
Swartkrans
35 ca.
Ivromdraai
3
East Africa
Olduvai Gorge
3
Kanam
1 (?)
Garusi
1
Sahara
Tchad
1
Palestine
Tell Ubeidiya
1
Java
Djetis Beds
2
China
drugstore
1 (?)
The order in which these sites have been arranged in Table 4 is
based primarily on the amount of information available for each
specimen or group of specimens. It may also reflect relative age,
although this is not sure.
The South African Australopithecines:
Time, Space, and Taxonomy
The most numerous and most fully described of early homi-
nids are the Australopithecines of South Africa. They seem to be
divided into two successive populations of different sizes and
degrees of resemblance to Homo. What we know about them we
have learned as a result of the energy and devotion of a few dedi-
cated South African anatomists and paleontologists, notably Dart,
Broom, and Robinson. Their success is due in part to the fact that
in Africa south of the Sahara limestone is scarce. With the modern
building boom in Johannesburg and other cities along the Rand,
whatever deposits there are have been subjected to quarrying,
232
The Earliest Hominids
dynamiting, and conversion into cement. Like most limestone, the
Transvaal deposits contain fissures and caves, many of which are
filled with sandy breccia, useless to limeworkers. These breccias
are packed with animal bones, mostly those of ungulates, but
some are the bones of baboons and a few of hominids. Since 1924
when Dart first identified the infant skull of Australopithecus
africanus, fragmentary remains of many other hominid individ-
uals have been tediously cut out of the breccias.
So far only five such sites contain these remains. They are lo-
cated in three widely separated regions (see Map 5). Taung,
where the first find was made, is in Bechuanaland, six miles west
of Taung Station and eighty miles north of Kimberley. Kromdraai,
Sterkfontein, and Swartkrans are clustered together within a
space of three miles located six to nine miles north-northwest of
Krugersdorp in the Transvaal. Makapansgat is near Potgietersrust,
north Transvaal, 165 miles north of Pretoria. From Taung to the
three central sites is about 200 miles, and from the latter to Maka-
pansgat about 150. Taung is in dry country, whereas the other
four are well watered. These differences in relative humidity ex-
isted during the time of the Australopithecines, as they do today.
The Taung site consists of a dolomite plateau scored by deep
cracks. Into these crevices animal bones had fallen or been
washed, along with sand, which cemented them into two succes-
sive breccias, a gray below and a pink above. The Sterkfontein site
is a cave which had a hole in its roof in Australopithecine time. As
the bones inside had fallen through the roof, it could not have
been a habitation site. The other three, Makapansgat, Swartkrans,
and Kromdraai, were apparently ordinary caves.4
The remains from all these sites are as fragmentary as the bony
refuse from a lamb stew, and consist largely of teeth. It is impos-
sible, therefore, to say how many individuals are represented.
However, since the initial discovery at Taung, in 1924, of an in-
fant skull, which was promptly named Australopithecus africanus,
the remains of several hundred similar hominids have been col-
lected. This skull came from the pink breccia. Following hoary
paleontological tradition the hominids were initially classified
into four genera and six species, as follows: A. africanus, Taung;
4 G. B. Barbour: “Ape or Man?” OJS, Vol. 49 (1949), pp. 129-45.
The South African Australopitliecines
233
Plesianthropus transvaalensis, Sterkfontein; A. prometheus, Ma-
kapansgat; Paranthropus crassidens, Swartkrans; Telanthropus
capensis, Swartkrans; and Paranthropus robustus, Kromdraai.
Dart named the two species of Australopithecus, Broom and Rob-
inson the others. The commonly known subfamily named Aus-
tralopithecinae, which encompasses all of them, can be used in-
formally in the guise of Australopithecine.
The succession of these five sites has been determined by three
234
The Earliest Hominids
methods: soil analysis from the breccias, faunal association, and
tool association. Soil analysis, which Brain conducted,5 is used to
determine whether a climate was wet or dry and what pattern of
climatic changes occurred while each breccia was forming. In both
Sterkfontein and Swartkrans the climate started out like that of
today, grew drier, and then again wetter; but at Swartkrans the
dry interval was much less intense than at Sterkfontein, and the
pattern of change is different. As the two caves lie near each other
in the same valley, this seems to indicate that two different cli-
matic cycles are involved. Kromdraai, also nearby, had a wetter
climate than today; it grew a little less wet as time went on. The
other two sites could not be studied in this way.
The faunal study conducted by Miss Ewer6 indicates that
Taung, Sterkfontein, and Makapansgat were roughly contempo-
raneous, or at least overlapping in time, and that Swartkrans came
later, after a gap. The fauna assigned to Kromdraai came from a
separate site 100 yards away from the Australopithecine-bearing
cave. It is even later than that of Swartkrans and could have been
Early Middle Pleistocene.
In comparing the South Africa cave faunas as a whole with those
of East Africa, Miss Ewer found that they are as old as Omo or
older. They contain 47 per cent living genera and 12 per cent liv-
ing species. Omo has 60 per cent living genera and 21 per cent liv-
ing species. These results imply that the South African Austra-
lopithecines appeared on the local scene as early as did their
counterparts farther north.
The archaeological evidence that constitutes the third method,
tool association, has nothing to do with the question whether or
not the Australopithecines made or used tools, which will be dealt
with later. It is concerned only with associations outside the caves
themselves. In the terraces of the nearby Vaal River Valley archae-
ologists, working independently of the fossil-hunters, have dis-
covered a three-stage tool sequence, starting with Early Oldowan.
5C. K. Brain: “New Evidence for the Correlation of the Transvaal Ape-Man-
Bearing Cave Deposits,” TCPC ( 1957), pp. 143-8.
Brain: “The Transvaal Ape-Man-Bearing Cave Deposits,” TMM, No. 11
(1958).
B. E. Sabels : review of Brain’s work, AJPA, Vol. 17, No. 3 (1959), pp. 247-9.
6 Ewer: op. cit.
The South African Australopithecines 235
By comparing the climates of the caves with those of the succes-
sive valley levels, Oakley 7 has found that the three early sites,
Taung, Sterkfontein, and Makapansgat, were probably con-
temporaneous with the Early Oldowan level, whereas the other
two, particularly Kromdraai, belong with the fully evolved Oldo-
wan tool level, which is geologically separate from the first.
Although each of the three methods — soil analysis, faunal as-
sociation, and tool association — has its limitations, the cumulative
effect of the three is impressive. The early sites are thus associated
with Omo and Kanam, Swartkrans with Olduvai Bed I, and
Kromdraai could even have overlapped the beginning of the Mid-
dle Pleistocene.
This division of the five sites into two consecutive, nonover-
lapping groups agrees with the anatomical evidence. The first
three caves contained small hominids about the size of a living
human Pygmy or even smaller, under five feet or 150 cm. tall and
weighing less than 100 pounds. The Swartkrans and Kromdraai
creatures were taller and heavier, within the full-sized human
range in both dimensions. In fact Swartkrans may have weighed
as much as 150 pounds.8 There is some question about Telanthro-
pus’s size and status; that will be discussed later. That we are
dealing with samples of two successive populations makes good
sense ecologically. It is doubtful that two closely related species
or subspecies of hominid could both survive competition for the
same food supply while living within a mile of each other in the
same valley. Whether the big species evolved out of the small one,
or simply replaced it after invading from the north, this evidence
does not tell us.
Despite the proliferation of taxonomic names originally given
these hominids, the group as a whole was no more variable than
the living chimpanzee, including its pygmy form. Washburn,9 sec-
onded by Dart himself,1 has proposed that all the Australopithe-
7 K. P. Oakley: “Dating the Australopithecines,” TPCP (1957), pp. 155-7.
8 These estimates, based principally on pelvis size, were made by W. L. Straus,
Jr. See S. M. Gam and A. B. Lewis: “Tooth Size, Body Size, and ‘Giant’ Fossil
Man,” AA, Vol. 60, No. 5 (1958), pp. 874-80.
9 S. L. Washburn, in discussion of E. Mayr: Taxonomic Categories in Fossil
Hominids (Cold Spring Harbor Symposia, Vol. 15, 1950), p. 118.
1 R. A. Dart: “Australopithecus prometheus and Telanthropus capensis,” AJPA,
Vol. 13, No. 1 (1955), pp. 67-96.
236
The Earliest Hominids
cines so far known constitute a single genus, and Oakley 2 has
further proposed that this genus can be divided into no more than
two species, Australopithecus africanus for the Lower Pleistocene
specimens and A. robustus for the later ones.
We now have three sets of names for the South African Australo-
pithecines: five site names, four of which refer to separate and
single kinds of animal each, and only one of which houses two
kinds; five generic names as originally proposed, two of which
encompass two species each; and one new genus with two species
separated by a time threshold. Of the three sets the only one
which cannot be changed is the first. I shall therefore follow the
current procedure employed by experts in this field and call the
specimens by the names of their sites, except for Telanthropus,
whose status is in doubt anyhow. The list of names is now: Taung
(A. africanus ); Sterkfontein ( Plesianthropus ); Makapansgat (A.
prometheus ); Swartkrans ( Paranthropus crasshlens) ; Kromdraai
( P ■ robustus ) ; and Telanthropus ( Telanthropus).
The Australopithecine Cave Sites
Unlike Bed I of Olduvai, not one of the five Australopithecine
sites can be called with certainty a habitation or occupation site;
and since Taung and Sterkfontein are mere refuse pits, they are
out of the question. The breccias of all five were broken by quarry-
men into blocks, out of stratigraphic context. Even if Makapans-
gat, Swartkrans, and Kromdraai had contained superimposed oc-
cupation floors, we could not list them in order. Howell, in 1959,
considered that the way some of the ungulate long bones were
split, and their lack of tooth marks, indicated that the Australo-
pithecines might have brought the bones into the caves. But no
primate except man has been known to live in caves,3 and even
men will rarely enter them unless they have fire, or at least a light.
Caves are dark, and hominids do not have night vision. Caves are
dank and clammy; without fire they are uncomfortable. Caves also
2 Oakley: “Dating of the Australopithecines of Africa,” AJPA, Vol. 12, No. 1
(i954), PP- 9-23-
8 A possible exception is the crab-eating macaque of the Philippines, which
lives along the shore and has no predators to fear but man.
Did the Australopithecines Make Tools? 237
harbor predatory beasts, like tigers, and smaller but almost equally
objectionable porcupines. We have no evidence that human be-
ings lived in caves before they had fire. And none of the Austra-
lopithecines had it. ♦
In the Limeworks Cave at Makapansgat a count was made of
the animal bones removed from the breccia. Ninety-two per cent
were the bones of antelope of different species and sizes. Ba-
boons accounted for 1.7 per cent, and the Australopithecines for
only .26 per cent. Had the Australopithecines been both residents
and cannibals, like Sinanthropus, the count of their own bones
would be higher. Whoever or whatever animal it was that lived
almost exclusively on antelopes must have been an accomplished
hunter, a far better one than the contemporary hominids of Oldu-
vai and Tell Ubeidiya (Israel), as we shall soon see.
Howell attributes to Desmond Clark a statement that these
caves contained springs and that animals entered them to drink.4
This may well be true. But the Australopithecines would not
dwell in a busy public watering place frequented by large preda-
tors as well as by their prey.
Did the Australopithecines Make Tools?
We do not know whether the Australopithecines made tools.
We only know that someone was flaking tools in Australopithecine
country when those hominids lived there, and that those tools
found their way into two successive terraces of the Vaal. If the
Australopithecines did not make the stone implements in question,
then they could only have been made by true men, of whom no
physical trace has yet been found.
Yet true men could hardly have coexisted with Australopithe-
cines in a single valley for over a hundred thousand years ( a mini-
mal estimate ) without having exterminated their close rivals for
the food supply. We are left with the circumstantial evidence that
the Australopithecines probably did indeed make the stone tools.
There is no reason at all for stone tools, or any other kind of im-
plements, to be found in the breccias except by coincidence. The
4 Howell: “The Villafranchian. . . .”
238
The Earliest Hominids
Australopithecines almost certainly did not live in the caves, and
a dead hominid who is being dragged into a cave by carnivores, in
several pieces, is not likely to bring his tools with him.
At Makapansgat seventeen “pebble- tools” have been found in
breccia above the Australopithecine-bearing layer.5 At Kromdraai
two pieces of intrusive rock, not identified as implements, have
also been found.6 At Sterkfontein worked pebbles have been re-
moved from an upper level of breccia which also contained an
Australopithecine maxilla and several teeth.7 Leakey calls this
level Middle Pleistocene. A direct bit of evidence is Schepers’s dis-
covery of a piece of “flint-like rock” imbedded in the skull and en-
docranial cast of a Kromdraai specimen, who may well have died
as a result,8 but the victim could have lived late enough to have
been killed by a pioneering Middle Pleistocene Homo. Schepers
was obliged to destroy this object in cleaning the matrix.
Dart had, in 1948, a collection of fifty-eight baboon skulls from
Taung, Sterkfontein, and Makapansgat, forty-two of which
showed depressed fractures, some in the form of double dents as
if made by blows from the distal condyles of an ungulate femur.
More had been struck on the left side than on the right. He states
that these baboons had been tapped on the head by such a bone
weapon, held in some kind of hominid’s hand, in this case, the
hand of an Australopithecine.9 In recent years he has had pub-
lished widely his theory that some of the bones, horns, and teeth
recovered from the breccias were used by the Australopithecines
as weapons and tools.1 I have handled some of Dart’s specimens
5 Brain, C. van R. Lowe, and Dart: “Kafuan Stone Artifacts in the Post-
Australopithecine Breccia at Makapansgat,” Nature, Vol. 175, No. 4444 (1955),
p. 16-18.
6 L. S. B. Leakey: “A New Fossil Skull from Oldoway,” Nature, Vol. 184, No.
4685 (1959), pp. 491-3.
7 Oakley, in comment on paper of M. Bonnardel: “La Main et L’Outil,” in
Les Processus de L’Hominisation, A Delmas, ed. (Paris, 1958), p. 131.
8G. W. H. Schepers, “The Endocrinal Casts . . . ,” pp. 173-4. Also personal
communication.
9 Dart: “The Makapansgat Proto-Human Australopithecus Prometheus,” AJPA,
Vol. 6, No. 3 (1948), pp. 259-84.
Dart: “The Predatory Implemental Technique of Australopitheus,” AJPA, Vol.
7, No. 1 ( 1949), pp- 1-16.
1 Dart: “The Makapansgat Australopithecine Osteodontokeratic Culture,” PTPC,
1955, PP- 161-71.
Dart and J. W. hatching: “Bone Tools at the Kalkbank Middle Stone Age Site
Did the Australopithecines Make Tools? 239
and find them as unconvincing as his ingenious theory is unneces-
sary. Recently Pei and others have found the same kind of bone
“tools” as Dart’s in a Chinese mammalian fauna of Late Pleisto-
cene date, without any evidence of man’s presence.2
Whatever made them, the paired depressions on the baboon
skulls are the likeliest evidence we have that the Australopithe-
cines hunted at all, but they are not convincing, for two reasons.
(1) Only one of the three sites, Makapansgat, was an ordinary
cave into which predators dragged the remains of their kills. Most
of the bones were those of adult animals. Taung and Sterkfontein
were holes into which animals fell, or their bones were washed.
Neither was a habitation site. Both were naturally formed refuse
pits. (2) Of the twenty- two photographs of punctured baboon
skulls shown in Dart’s 1949 article, sixteen are adult, one is juve-
nile, three are infant, and two are of undetermined age. If 80 per
cent of Australopithecus’s primate victims were adult, then he was
a mature hunter, like the unidentified killer who left the bones of
his victims in Makapansgat cave. Whoever killed the antelopes of
Makapansgat and the baboons of all three sites was therefore a
much better hunter than the hominids who inhabited Olduvai
Gorge and the Jordan Valley, where nearly all the animals killed
were infants. It is hard to believe that the South African Austra-
lopithecines were better hunters than their relatives farther
north.
The Postcranial Skeletons of the South
African Australopithecines
A hominid can hunt only if his hands are free to hold weap-
ons, and his hands are free only if he can stand, walk, and run
bipedally. Whether or not an animal stood erect can be deter-
mined by studying the bones of its postcranial skeleton. In the
and the Makapansgat Australopithecine Locality, Central Transvaal, Part 2, The
Osteodontokeratic Contribution,” AB, Vol. 13, No. 51 (1958), pp. 94-116.
And many other titles.
2 W. C. Pei, W. P. Huang, C. L. Chiu, and H. Meng: “Discovery of Quater-
nary Mammalian Fauna at Ch’ao-tsun, Chien-An County, Hopei Province,” VP,
Vol, 2, No. 4 (1958), pp. 226-9.
240
The Earliest Hominids
published literature on the Australopithecines 3 we have informa-
tion on twelve such bones, or sets of bones, as shown on Table 5.
To this list may be added the following pieces of an adult fe-
male skeleton from Sterkfontein, which has not yet been fully de-
scribed : eight thoracic vertebrae, six lumbar vertebrae, a sacrum,
two nearly complete pelvic bones, and a piece of femur, as well as
the body of a lumbar vertebra from another individual.4
Combining the above with the list on Table 5, we find that
Taung has no postcranial bones, Sterkfontein twenty-five, Maka-
pansgat five, Swartkrans two, Kromdraai eight, and Telanthropus
one. As some of the bones came in groups from single individuals,
the entire set could represent as few as seven or eight creatures.
The Sterkfontein Vertebrae and Ribs
The adult female found by Robinson at Sterkfontein
had eight of a putative twelve thoracic vertebrae, which have not
been described, and six lumbar vertebrae. This number is interest-
3 The bibliography is exhaustive. Here are some of the basic references in
which new specimens are reported.
R. Broom and Schepers: “The South African Fossil Ape-Men, the Australo-
pithecinae,” TMM, No. 2 (1946).
Broom, J. T. Robinson, and Schepers: “Sterkfontein Ape-Man, Plesianthropus ”
TMM, No. 4 (1950).
Broom and Robinson: “Swartkrans Ape-Man,” TMM, No. 6 (1952).
Robinson: “Telanthropus and its Phylogenetic Significance,” AJPA, Vol. 11,
No. 4 (1953), pp. 445-501.
Robinson: “The Dentition of the Australopithecinae,” TMM, No. 9 (1956).
Dart: “ Australopithecus africanus, the Man-ape of South Africa,” Nature, Vol.
115, No. 2884 (1925), pp. 195-9.
Dart: “The Makapansgat Proto-human Australopithecus prometheus,” AJPA,
Vol. 6, No. 3 (1948), pp. 259-84-
Dart: “The Adolescent Mandible of Australopithecus prometheus,” AJPA, Vol.
6, No. 4 (1948), pp. 391-412.
Dart: “The Cranio-facial Fragments of Australopithecus prometheus,” AJPA,
Vol. 7, No. 2 (1949), pp. 187-214.
Dart: “Innominate Fragments of Australopithecus prometheus,” AJPA, Vol. 7,
No. 3 (1949), PP- 301-38.
Dart: The Second or Adult Female Mandible of Australopithecus prome-
theus,” AJPA, Vol. 12, No. 3 (1954), pp. 313-43-
Dart: “ Australopithecus prometheus and Australopithecus capensis,” AJPA,
Vol. 13, No. 1 (1955), pp. 67-96.
Dart: The Second Adolescent (Female) Ilium of Australopithecus prome-
theus,” AJPA, Vol. 2, No. 1 (1957), pp. 73-82.
4 Robinson: “The Dentition of the Australopithecinae,” pp. 161, 170.
241
The Pelvis of Australopithecus
TABLE 5
AUSTRALOPITHECINE POSTCRANIAL
BONES
Bone
Common Name
Site, Animal
No. of Bones
Os Coxae
Pelvic bone
Makapansgat
2
Sterkfontein
2
Swartkrans
1
Femur
Thighbone
Makapansgat
1
Sterkfontein
1
Talus
Ankle Bone
Kromdraai
1
Phalanges (Foot)
Toe Bones
Kromdraai
2
Scapula
Shoulder Blade
Sterkfontein
1
Clavicle
Collarbone
Makapansgat
1
Humerus
Upper Arm Bone
Makapansgat
1
Sterkfontein
1
Kromdraai
1
Radius
Outer Lower Arm Bone
Makapansgat
1
Telanthropus
1
Ulna
Inner Lower Arm Bone
Kromdraai
1
Capitatum
A Wrist Bone
Sterkfontein
1
Metacarpals
Hand Bones
Kromdraai
1
Swartkrans
1
Phalanges (Hand)
Finger Bones
Kromdraai
1
ing since only about 5 per cent of human skeletons have six; 92 per
cent have five, and 3 per cent have four. The macaques usually
have seven, the chimpanzees four, and the gorillas four or three.
According to Robinson, when the lumbar vertebrae are articu-
lated with the sacrum a distinct lumbar curve is visible, as in man.
If and when Robinson’s statement is substantiated, we shall have
the best evidence yet that the Sterkfontein Australopithecines
stood and walked erect.
None of the ribs found at this site have yet been described.
The Pelvis of Australopithecus
No single bone is more sensitive to changes of posture
and locomotion than the pelvic bone, or os coxae, which is com-
posed of three elements fused together, the ilium, ischium, and
pubis. Of the four pelvic bones that have been described in pub-
lications, the two from Makapansgat 5 are immature and the ones
from Sterkfontein and Swartkrans are adult.6
In comparing these bones to the pelves of man and other pri-
6 Dart: “Innominate Fragments . . “The Second Adolescent. . . .”
6 Broom et al: “Sterkfontein . . "Swartkrans Ape-Man.”
242
The Earliest Hominids
mates we are severely limited because we have no pelvic bones for
Proconsul or other Dryopithecines, or for fossil men older than the
Neanderthals. We do have the bones of modern men, however,
and of apes and Old World monkeys. All four specimens resemble
those of living men in general form and in most details, but they
differ from the human pattern in a few features.7
Fig. 20 The Anatomy of the Human Pelvic Bone ( Os Coxae ) .
For example, in man the outer surface of the ilium is ordinarily
divided into three planes, one each for the attachment of the
gluteus maximus, gluteus medius, and gluteus minimus muscles,
and these planes are set at different angles. In neither Australo-
pithecines nor apes are these planes differentiated. However,
Dart ( 1957 ) has found the same condition in the pelvis of a mod-
7 L. W. Mednick: “The Evolution of the Human Ilium,” AJPA, Vol. 13, No. 2
(1955), PP- 203-216.
W. E. LeG. Clark: “The Os Innominatum of the Recent Ponginae, with Spe-
cial Reference to That of the Australopithecinae,” AJPA, Vol. 13, No. 1 (1955),
pp. 19-28.
Dart: “The Second Adolescent . . . ,” pp. 73-82.
The Pelvis of Australopithecus 243
ern Pygmy. His discovery implies that in small and light bipedal
animals, like Pygmies, Sterkfontein, and Makapansgat (if the two
latter were bipedal ) , ridged areas of attachment for these muscles
are unnecessary; but Swartkrans was as heavy as a full-sized man.
Also the ischial bone differs both among the Australopithecines
and between them and man. This is the lowest and rearmost of the
three fused pelvic bones. As stated in Chapters 4 and 5, in the
Old World monkeys and apes the ischium extends far down and
ILIUM
CHIMPANZEE AUSTRALOPITHECUS MODERN MAN
(SWARTKRANS)
Fig. 21 Pelvic Bones of Ape, Australopithecus, and Man. Note the differences
between these three Hominoids in the ilium and in the ischium. Whereas the
Australopithecine bone is generally short and compressed, as in man, the
tuberosity of the ischium is set well below the level of the acetabulum, and in this
feature it is intermediate between man and ape. (Drawings after Abbie, 1961.)
to the rear, and its tuberosity flares outward. In man it is short
and bends inward. Only in man does the gluteus maxitnus muscle
cover the ischial tuberosity. Makapansgat’s ischium is the most
manlike, Sterkfontein’s slightly less so, and Swartkrans’s is nearly
apelike.
This evidence suggests that none of the Australopithecines sat
exactly as man does, and that Sterkfontein and Makapansgat
were more nearly human in posture than Swartkrans. LeGros
Clark, Robinson, and some other anatomists agree that the Austra-
lopithecines stood erect, but also that their posture was less per-
fect than ours, and Robinson ( 1956) suggests that the earlier spe-
cies ( Australopithecus africanus ) and the latter one (A. robus-
tus) were members of separate lines which had acquired the
erect posture, such as it was, independently. Washburn has re-
244
The Earliest Hominids
cently suggested that the Australopithecines in general could not
have walked erect as we do, but could only run while erect,
dropping down on all fours when at rest.8
The undescribed sacrum mentioned by Robinson (1954) is
nearly but not wholly complete. It is said to be short and broad,
and to bear a close general resemblance to those of modern men.
Of the pelvic bones (os coxae) found with the sacrum Robinson
states that they are oriented in the human fashion and that they
include a pubic symphysis (juncture of the two pubic bones), de-
scribed as short. In the apes the pubic symphysis is longer than
in man. Robinson concludes that the adult female whose mid-
portions he so painstakingly removed from the breccia was small,
weighing only 40 to 50 pounds, but that her pelvis was broad and
sturdy, fully adequate to support her weight in bipedal locomo-
tion.
The various reconstructions and conclusions quoted above,
variable as they are, are of considerable theoretical interest in the
study of human locomotion. They offer the possibility that more
than one evolutionary line within the Hominidae could have ac-
quired the erect posture independently, just as more than one
reptile became a mammal, and they suggest a reason why stand-
ing and walking erect became advantageous in the first place. If
piimates like the earlier Australopithecines, and the forerunners
of Homo, were able to walk erect, they could have used their
hands to make and carry tools. Rut if they moved about on all
fours except when dashing after game, they could not have car-
ried tools, even if they made tools when sitting down. Unless they
could walk, they could not have carried water and food from
source to camp. Much depends on their means of locomotion,
which can be further elucidated by studying other bones, particu-
larly those of the legs and feet.
The Legs and Feet of Australopithecus
Although no whole femur has been described, we have de-
scriptions of two femoral heads, only one of which is complete
8 Washburn: “Tools and Human Evolution,” SA, Vol. 203, No. 3 (i960) pp
63-75-
The Legs and Feet of Australopithecus 245
enough to be useful,9 and of one distal, or lower, end.1 The nearly
complete head belonged to a specimen from Sterkfontein. To it is
attached the crushed shaft of most of the rest of the femur, which
Broom estimated to be 310 mm. long, barely within the lower
border of the human range. The position of the head on the
shaft fits that of the acetabulum of the pelvis described above.
Although it faces outward, it does not rotate as far forward as in
man, nor are its areas of muscle attachment as strongly devel-
FEMUR
CHIMPANZEE
Fig. 22 The Distal End of the Femur in Australopithecus, Ape, and Man. In
the Sterkfontein femur, the angle between the shaft and the condyles is greater
than in either chimpanzee or man. In the end view (bottom row), the Sterkfontein
condylar head is squarer and more deeply notched than in either chimpanzee
or man. ( Drawings after Broom, 1946. )
oped." All that can be said of the second specimen, the one from
Makapansgat (Bone, 1955) is that it is believed to come from a
different individual than did the other bones from that site, and
that it is larger than the Sterkfontein specimen.
9 Broom et al.: “Sterkfontein Ape-Man, Plesianthropus.”
1 Broom: The Occurrence and General Structure of the South African Ape-
Men, in Broom and Schepers: “The South American Fossil. . . .”
Broom and Robinson: Further Evidence of the Structure of the Sterkfontein
Ape-Man, Plesianthropus, Part I, of Broom et al.: “Sterkfontein Ape-Man. . . .”
E. Bone: “Quatre Fragments Post-Craniens du Gisement a Australopitheques de
Makapansgat (N. Transvaal),” L’Anth., Vol. 59, No. 5/6 (1955), pp. 462-9.
Bone and Dart: “A Catalogue of Australopithecine Fossils Found at the Lime-
works, Makapansgat,” A/PA, Vol. 13, No. 4 (1955), pp. 621-4.
H. M. Kern, Jr., and Straus, Jr.: “The Femur of Plesianthropus transvaalensis,”
AJPA, Vol. 7, No. 1 (1949), pp. 53-78.
" '^le great trochanter is less sharply lipped than in a Bushman femoral head
used for comparison; the trochanteric fossa is shallower, and there is no well-
developed trochanteric crest.
The Legs and Feet of Australopithecus 247
of its complicated form, the degree of rigidity or mobility of the
ankle joint. In the apes the advantage lies in mobility, for the foot
is used more or less as a hand. In bipedal primates rigidity is ad-
vantageous, for, while the animal is walking, each talus bears in
turn the entire weight of the body. In the quadrupedal monkeys,
whose gait places less strain on this bone, an intermediate condi-
tion is more suitable.
Fig. 24 The Astragulus of the Australopithecines and Other Primates.
A. Homo sapiens, a woman; B. Kromdraai; C. Olduvai Child; D. Baboon;
E. Chimpanzee; F. Proconsul nyanzae. In the angle of the neck of the
astragulus to its main axis, the Australopithecines are intermediate between the
apes and baboons on the one hand, and men on the other. ( Drawings A, B, and D
after Broom, 1946; C after Leakey’s photograph, 1961; E and F after LeGros Clark
and Leakey, 1947. )
Seen from above, the Kromdraai talus may be compared with
those of several other primates, including that found with the
Olduvai child, which will be discussed later. Although it is broken
in the rear, enough is left so that the shape and general dimen-
sions can be reconstructed with some accuracy. It falls within the
human range in size, and probably in the length-breadth ratio,
and its neck and head point inward at an angle of about 26°, a
The Earliest Hominids
248
figure on the outer edge of the human range ( 18-25° ) and below
that of living apes ( 30-36° ) 3 and of Proconsul nyanzae ( ca.
35° )• In the form of its facets it appears both to be of the proper
size and to have the rigidity necessary for bipedal walking. It is
essentially human.
The two toe bones of the same animal are an incomplete first,
or proximal, phalanx of the left little toe, and a last, or distal,
phalanx of either the second or the third toe. Both bones appear
longer and a little more fingerlike than the corresponding bones of
most human feet.
All in all, the known leg and foot bones of the South African
Australopithecines are, like their pelvic bones, essentially manlike
and where they differ from the corresponding human bones they
are not particularly apelike. In some respects they point rather to
Proconsul and the baboons.
Fig. 25 The Foot Bones Found with the Olduvai Child (compared with
those of a gorilla and a Bushman ) . The drawing of the foot bones found with the
Olduvai child, minus the phalanges, which were not found, is tentative and subject
to revision when the original bones are illustrated in a scientific monograph. The
calcaneum is especially dubious because it is broken. So is the form of the
cuboid (among other bones), which is out of place in the available photographs.
Nevertheless, the Olduvai foot is clearly intermediate in many respects be-
tween those of ape and man. The heel part of the calcaneum lies on the outer
side of the foot, as in the ape, rather than under the center of gravity, as in man.
The ratio in length between the tarsal and the metatarsal bones is intermediate be-
tween those of ape and man, and the first metatarsal (great toe) projects some-
what forward and inward, in an intermediate fashion. In the Olduvai foot the fifth
metatarsal overlapped the fourth, pathologically. ( Drawing of gorilla after Raven,
1950; Olduvai foot after photographs in the National Geographic Magazine, i960’
and the Illustrated London News, 1961; Bushman after Keith and McCown, 1938.)
J R. Martin and K. Sailer: Lehrbuch der Anthropologie, Third Edition (Stutt-
gart: G. Fischer; 1958), Section 7, pp. 1117-18.
The Shoulder Girdle of Australopithecus
249
The Shoulder Girdle of Australopithecus
The primate shoulder girdle consists of two pairs of bones, the
scapulae or shoulder blades and the clavicles or collarbones.
These bones attach the arms to the trunk, not rigidly as the pelvis
is fastened to the sacrum, but flexibly, through the agency of mus-
cles and tendons, and remotely, by the articulation of the clavi-
cles to the sternum, the sternum to the ribs, and the ribs to the
vertebrae. As we have seen in comparisons between monkeys and
apes, the shape and position of the scapulae and clavicles sensi-
tively reflect each animal’s mode of locomotion.
SCAPULAR NOTCH
ORANGUTAN CERCOPITHECUS (VALLOIS)
(VALLOIS, L'OMOPLATE HUMAINE )
Fig. 26 The Austbalopithecine Scapula.
In the Australopithecine material one scapula from Sterkfon-
tein and one clavicle from Makapansgat 4 have so far been de-
scribed. Both are fragmentary.
4 Broom and Robinson: “Further Evidence of the Structure . . . ,” pp. 55-7.
Bone: Une Clavicule et un Nouveau Fragment Mandibulaire d’ Australopithe-
cus prometheus,” PAf, Vol. 3 (1955), pp. 87-101.
250
The Earliest Hominids
About half the Sterkfontein scapula is available, and luckily it is
the more interesting half, containing the part that meets the hu-
merus (upper arm bone) at the shoulder. Because ground mon-
keys, apes, and men all hold their arms in different positions, this
part of the scapula is a highly diagnostic portion of the primate
anatomy.
The Sterkfontein fragment has a long spine, as do apes and
men but not monkeys. The head of its spine projects far out over
the head of the humerus, as it does in apes and men but not in
monkeys. But the head is thick, as in man alone. Oddly enough,
the spine of this scapula is thin and straight, more like those of
Negroes than like those of any other modern race. What this
means I do not know.
In other respects the human image fades. Some details of the
upper and lower borders of the scapula and of the glenoid cavity
the concave facet on which the head of the humerus moves —
resemble those of apes and monkeys, especially the former, and
also suggest that the Australopithecines put more muscular strain
on their arms and hands than most living men do. In general, this
scapula resembles that of man as far as the position of the arm in
the shoulder is concerned, but where it differs from man’s it is
apelike rather than monkeylike.5
5 The Sterkfontein specimen has no scapular notch, which in man is usually a
deep, semicircular cavity at the base of the coracoid process, enclosing, with the
help of a ligament, the suprascapular nerve. This notch is usually present in the
Cebidae and man, and absent in prosimians, Old World monkeys, and apes, al-
though a trace of it turns up now and then in baboons. In man it is sometimes
absent altogether, as in 5 per cent of Pygmies, or it may take many variant forms,
including a bridged hole.
The lower border of the scapula also is not human in form. The so-called
axilospinal angle, the angle between the axis of the spine and that of its lower,
outer border (next to the axilla, or armpit) is 22 0 in Sterkfontein. The range for
apes is from 8° in the gibbon to 180 in the orang. An arboreal Old World monkey,
Cercopithecus sp., has an angle of 30°, and the individual human range is 24
to 58“. S
Another diagnostic angle is that formed between the axis of the glenoid cavity
and that of the scapular spine. Although this angle could not be measured in the
conventional way because of breakage, a substitute technique, which can be ap-
plied to comparable fragments of other scapulae, gives Sterkfontein an angle of
103°, wider than those for man (about 90°) and the living apes (85° and lower).
Broom, who was able to examine the specimen closely, said that the glenoid
cavity has a much larger attachment for the biceps muscle than does the corre-
sponding part in man. This indicates further that the Australopithecus scapula
251
The Arms and Hands of Australopithecus
The Sterkfontein clavicle (collarbone) is even less complete
than the scapula from the same site. Owing to its fragmentary
condition, it is useless in our present study.
The Arms and Hands of Australopithecus
We have three pieces of humerus : one almost complete bone
from Sterkfontein which is crushed flat except for the head; 6 a sec-
tion of shaft about 64 mm. long from Makapansgat; 7 and a distal
(elbow) portion from Kromdraai.8 These three specimens are
similar enough to be treated as a unit, if we keep in mind that
the first two came from smaller animals than the third.
The first bone was originally about 30 cm. long, within the
ranges for man and chimpanzee. A comparison of this figure with
the length of a femur from the same deposit, but not necessarily
from the same individual, gives a skeletal upper-arm— thigh in-
dex of 96. In apes the humeri are always longer than the femora,
so that the individual humero-femoral indices range from 111 to
147. In the Old World monkeys the index is always below 100,
and in man it is about 71 to 76. This evidence suggests that Sterk-
fontein’s upper arms were shorter than his thighs and that he
probably walked erect. At least, he did not walk on all fours the
way that apes do when on the ground.
The piece of shaft from Makapansgat is heavy, thick, ridged
with muscular crests, and structurally humanlike. It is more robust
than either the head just described or the elbow piece about to be
described.
The latter, from Kromdraai, is particularly significant. It in-
in question could not be mistaken for that of man, ape, or monkey, and that in
general it tends rather toward the apelike than the human.
H. V. Vallois: “L’Omoplate Humaine,” BMSA varia 1928-46 (Paris: Masson
et Cie; (1946).
Martin and Sailer: op cit., Vol. 1 ( 1957) p. 531.
G Broom and Robinson: “Further Evidence of the Structure . . . ,” p. 57.
7 Bone: “Quatre Fragments Postcraniens du Gisement de Makapansgat, N.
Transvaal,” L’Anth, Vol. 59, Nos. 5-6 (1955), pp. 462-9.
8 Broom: “The Occurrence and General Structure . . . pp. 114-15.
Straus, Jr.: “The Humerus of Paranthropus robustus,” AJPA, Vol. 6, No. 3
(1948), pp. 285-312.
252
The Earliest Hominids
eludes unbroken articular surfaces that permit and delimit the
movements of the forearm at the elbow, and are thus concerned
with both locomotion and the rotation of the hand when it is
free. In morphological details 9 this piece of shaft falls into the
same general category as do those of Proconsul, apes, and men and
is essentially different from the distal humeri of the Old World
monkeys, both arboreal and ground-living. In a few respects it is
unique. On the whole it suggests a degree of forearm mobility in-
termediate between those of apes and men, without settling, in
itself, the question of locomotion.
We have fragments of two real or alleged radii, the proximal
and distal ends of what is believed to be a single bone, from
Makapansgat, and an alleged distal extremity of Telanthropus.1
The first is doubtful. The head could be hominid or even pongid,
but the end piece is not necessarily that of a primate, for it resem-
bles the radii of carnivores, particularly cats. The second or Telan-
thropus piece, according to Robinson, can be matched among
modern human radii.
At Kromdraai Broom 2 found a fragment of ulna from the same
elbow joint as the distal piece of humerus described above. It is
only 3.7 cm. long, but it contains both the diagnostic articular
surfaces and the olecranon process — the hook that locks the bone
to the humerus when the arm is fully extended. Although it seems
human in most respects, it has two anomalies. One is a thin, hol-
low-backed ridge on the inner side of the olecranon process, the
other a deep hollow near the extremity of the articular surface.
9 As in Proconsul africanus, the apes, and man, the angle between the axis of
the shaft and that of the condyles is ioo° or more; in the Old World monkeys it
is about 90°. Also, the articulation with the radius is the same as in man, but the
articular surface for contact with the ulna is oval rather than round, unlike that
in either man or ape. The inner condyle is small and more pointed than it is in
either man or ape, and the outer condyle also is “not quite human” (Broom,
1950, p. 115). The supinator ridge on the lateral condyle, for the attachment of
the supinator muscle — a muscle joining the humerus and radius and concerned
with the rotation of the lower arm — struck Broom as being more apelike than hu-
man. He also found a deep furrow and pit between the same condyle and the
radial articular surface which is present neither in apes nor in men.
1Bone: “Quatre Fragments. . . .”
Robinson: “Telanthropus and its Phylogenetic Significance.”
2 Broom: “Paranthropus robustus Broom,” in Broom and Schepers: “The South
African . . . ,” p. 115.
253
The Arms and Hands of Australopithecus
Broom was able to find similar characteristics in about 1 per cent
of the human ulnae of various races, but he failed to find them in
ape ulnae.
The only carpal bone we have is a right os capitatum, or capi-
tate bone, from Sterkfontein.3 This is a small, more or less cubical
bone in the middle of the outer row of wrist bones, one which ar-
ticulates, in man, with five or sometimes six other bones, thus
forming the center of the wrist. Human in size and intermediate
between the corresponding bones of apes and men in mobility, it
is particularly human in its implication of a large and mobile
index finger.4
The metacarpal of one thumb, that from Swartkrans, throws
light on the possible manual activities of the larger Australopithe-
cines.5 It is 36 mm. long, which places it in the lower part of the
human range, and stoutly built. As seen from the side, it appears
to be more curved than the metacarpal of either men or apes
(Fig. 27). On the inner side of the rear or proximal end is a sharp
beak that once separated a pair of large sesamoid bones.
Sesamoids, so called because they look like sesame seeds, are
small bones which form in tendons over joints that are frequently
flexed in a single direction and with considerable force; they serve
3 Broom: “Plesianthropus transvaalensis Broom,” in Broom and Schepers: op.
cit.
W. E. LeG. Clark: “Observations on the Anatomy of the Fossil Australopithe-
cinae,” JANAT, Vol. 81, No. 3 ( 1947). Reprinted in Yearbook of Physical Anthro-
pology (New York, 1947), pp. 143-77.
4 In man the facets by which it articulates with the other bones around it are
broad and extensive, permitting considerable mobility. In the apes, particularly
the gorilla and chimpanzee, these facets are small. In the apes, however, areas
for the attachment of ligaments are extensive; in man they are smaller. In the
apes, which have long, slender hands, the capitate is long; in man it is shorter.
In the apes the bone is larger than in man.
The Sterkfontein capitate is of human rather than apelike size, being com-
parable to those of baboons and Bushmen. (Maximum lengths: Sterkfontein, 18.3
mm.; Bushman female, 18.3; Bushman male, 20.9; baboon male, 19.0; Proconsul
africanus, 23.0; gorilla male, 26.5; chimpanzee male, 26.5; orang male, 31.7 mm.)
It is relatively broad, like a man’s. In the development of facets and ligamental
attachments, it is intermediate between those of apes and those of men, but
closer to the human. In particular, it has a larger articulation for the head of the
second metacarpal, on the thumb side, than is found in the apes; in the gorilla
there is characteristically none at all.
5 Broom and Robinson: “Thumb of Swartkrans Ape-Man,” Nature, Vol. 164,
No. 4176 (1949), pp- 841-2.
254
The Earliest Hominids
as fulcrums. The patella or kneecap is our largest sesamoid bone.
Such bones are not characteristic of the thumbs of apes or men.
In Swartkrans their presence suggests a specialization for some
kind of coarse work beyond the needs of hunters, such as digging
for roots.
A metacarpal from Kromdraai, probably that of the index finger,
is 70 mm. long, as long as that of a man with long hands. In most
respects it is human-looking, but two grooves on the palmar side of
A B
Fig. 27 The Australopithecine
Hand Bones. The Metacarpal Thumb
Bones of Australopithecus: A. Swart-
krans; B. Bushman; C. European.
Note that the Australopithecine bone
is curved and hooked at the base.
Proximal Phalanges of the Third or
Fourth Finger in Man, the Olduvai
Child, and the Gorilla: D. Man;
E. Olduvai Child; F. Gorilla. In man
the shafts of the phalanges are
rounded and narrow; in the gorilla
and other apes they are broad and
spatulate. In the shape of this bone
the Olduvai child resembles the apes.
( Drawings A and B after Broom and
Robinson, 1952.)
the distal knuckle betray the former presence of another pair of
sesamoid bones. These are rare or nonexistent in apes and normal
in baboons. They can occur in man, but seldom in pairs.
Two proximal phalanges of the same hand belong to the second
and fifth fingers. In the apes these phalanges are flattened on the
underside, and their edges are lipped; this represents an adapta-
tion for brachiation in a heavy animal, and reaches an extreme
form in the gorilla. It is not found in notable degree in Proconsul
africanus, the Old World monkeys, or man. The Kromdraai pha-
langes also lack it. No better indication is needed that this animal
was not a brachiator and that if his ancestors once had this adap-
tation they subsequently lost it.
The Skulls, Jaws, and Teeth of Australopithecus 255
Australopithecus, a Primate Mermaid or a Unique Hominid?
W e n o w have the impression that the South African Australo-
pithecines from the neck down were a kind of primate mermaid, a
somewhat composite animal. Either the Australopithecines were
going through a transitional stage of evolution that we have not
seen before, or the postcranial bones are a mixed bag of spare
parts of Australopithecines, apes, big baboons, and men. In gen-
eral, if not in every case, the first explanation is the more plausi-
ble.
No animal can be understood merely through comparison with
others, for each species is an entity in itself, an organism adapted
to its own kind of existence. From the study just made of the
postcranial bones of the South African Australopithecines it may
be concluded that these creatures must have followed a way of
life no longer seen on the earth. The resemblances of individual
bones to those of other primates living and extinct indicates prin-
cipally that these creatures drew on the same primate storehouse
of genetic potentialities as did monkeys, apes, and men. Moreover,
it is evident that the biological forces of parallelism, differential
evolutionary rates, and neoteny shaped a special organic whole to
fit a new ecological requirement in a creature whose ancestors had
parted company with those of the subfamily Ponginae before the
latter had become fully specialized.
As far as the postcranial skeleton is concerned, Homo could
have been descended from some kind of Australopithecine, more
like Australopithecus africanus, perhaps, than A. robustus. But we
can say no more until we have studied the rest of the skeletal
parts of the whole animals. The skulls and teeth will shed more
light on the relationships between Homo and his early South Afri-
can relatives.
The Skulls, Jaws, and Teeth of Australopithecus
Available published accounts G of the finds in the five caves
that have yielded the known remains of the South African Austra-
lopithecines include descriptions of the whole and fragmentary
6 The bibliography is essentially the same as that for the postcranial bones.
T he Earliest Hominids
256
brain cases, jaws, and teeth of at least twenty-one individuals or at
most about forty, three to seven times as many creatures as are ac-
counted for by the postcranial bones. Each of the South African
cave groups contains at least one skull, more or less complete.
However, there is none for T elanthropus, whose presence from the
neck up is indicated only by one nearly complete and one very
fragmentary lower jaw, and a piece of palate.
TABLE 6
SKULLS, JAWS, AND TEETH
OF AUSTRALOPITHECINES
Taung
1 skull, six years old, milk teeth
Makapansgat
3 fragmentary skulls
2 maxillae
3 mandibles
1 molar tooth
Sterkfontein
2 crania (one almost complete)
Kromdraai
1 cranium, fragmentary
2 mandibles
various teeth
Swartkrans
8 crania in various conditions
5 maxillae
10 mandibula
100 ca. loose teeth
Swartkrans,
Telanthropus
2 fragmentary mandibles
1 maxillary fragment
Before going further let us familiarize ourselves with some
technical terms that will appear in the descriptions of skulls and
their parts. The skull, technically, includes the entire structure:
the bones of the head, face, and both jaws. The cranium does not
include the lower jaw. A calvarium is a faceless and jawless brain
case; a calva consists of the top of the brain case only, without the
base; a maxilla, of the upper jawbone (one on each side); and a
mandible or mandibula, of the lower jawbone.
The Brain Case and Brain of Australopithecus
Next to the teeth, the brain case is the most human-looking
feature of this creature, as we can see (Fig. 28) from the pro-
The Brain Case and Brain of Australopithecus
257
Fig. 28 Skull Profiles of Australopithecines, Gorilla, and Baboon.
files of four skulls: the infant Taung, supposedly aged six years; an
infant from Swartkrans, supposedly aged seven years; an adult
male (?) from Sterkfontein; and an adult female (?) from Swart-
krans, greatly restored. In all of them the skull is long and rather
258
The Earliest Hominids
high, with a sloping forehead, and the brain case rises above the
brow ridges as high, in relation to its total size, as in some early
and primitive human crania. The foramen magnum is located
near the center of the skull base, as far from the rear as in some
human races; and the occipital condyles point downward as in
man instead of obliquely to the rear as in apes. In addition, the
occipital crest, which marks the upper limit of neck-muscle at-
tachments, is even lower than in some human skulls, and much
lower than in either ape or baboon. The brow ridges are heavy but
no more so than in some fossil and even a few modern human
skulls. As in man and only rarely in apes, the mastoid process
( a protuberance of the occipital bone ) is present, and it is conical
as in man. Since this anatomical landmark helps to anchor the
muscles that hold the skull erect on either side rather than from
behind, we may infer from the position of this process that the
Australopithecines had a human rather than an apelike neck. All
in all, the hominid skull form had been completely achieved by
Australopithecus, which suggests but does not prove that this
animal stood erect.
A human brain size, however, had not been achieved, nor even
approximated. The Taung child of six had a brain size of 494 cc.,7
which could have attained a maximum of 543 cc. had the creature
lived to maturity and had his brain continued to grow at the same
rate as in modern man, which cannot be assumed. The Maka-
pansgat brain case is known only from a single occipital bone. Its
capacity has been estimated, very speculatively, at 600 cc.8 Two
Sterkfontein skulls complete enough for accurate measurements
have capacities of 435 cc. and 480 cc. Thus, the brains of the
smaller and earlier Australopithecines range from a known 435 cc.
to a possible 543 cc. and a speculative 600 cc.
A quite fragmentary Kromdraai skull may possibly have at-
tained 650 cc., although this is unlikely; and the various Swart-
krans skulls, none of which is complete enough to measure prop-
erly, have been estimated at 700 cc. Washburn denies that any
of these skulls can have exceeded 550 cc.: the foramen for the in-
7 C. J. Connolly: External Morphology of the Primate Brain (Springfield, 111.:
Charles C Thomas; 1951). See pp. 293-5.
8 Clark: “The Os Innominatum. . . p. 122.
The Brain Case and Brain of Australopithecus 259
ternal carotid artery, one of the vessels that feeds blood to the
brain, is no larger than those of apes.9
According to the principle of allometry (see page 25), which
governs the differences in proportions of similar animals of differ-
ent sizes, if the large Australopithecines had the same kind of
brains as the small ones, the cubic capacity figure of the large ones
would be absolutely larger and proportionately smaller than those
of the small group. The difference between the two groups seems
to be about 100 cc., or about 20 per cent. However, the differ-
ence in body size may have been in the order of 50 to 100 per
cent. These figures do not suggest any increase in intelligence be-
tween Australopithecus africanus and Australopithecus robustus
such as one might expect to find if the second species were evolv-
ing from the first in the direction of man.
The variously estimated range in brain size of 435 to 700 cc.
ascribed to the Australopithecines by several authors falls within
the middle and upper parts of the anthropoid ape range, which is
325 to 685 cc.; and it may not exceed that of the largest known
gorilla — 685 cc. The brains of both putative species of Australo-
pithecines were a little larger for their body size than the brains of
the three living great apes, but not enough larger to indicate,
without supplementary evidence, a substantial difference in in-
telligence. It is extremely unlikely that they could speak.1
9 Washburn and V. Avis: “Evolution of Human Behavior,” in A. Roe and G. G.
Simpson. Behavior and Evolution (New Haven: Yale University Press; 1958)
P- 430.
Although an approximately equal amount of blood also flows up through the
vertebral arteries, this fact alone does not invalidate Washburn’s conclusion, be-
cause the ratio of blood contributed by each pair of arteries is the same in all
living primates.
1 Schepers, who studied the internal brain casts of the Taung infant, five
Sterkfontein skulls, and a fragmentary juvenile specimen from Kromdraai, after
having constructed elaborate charts of the surface configurations of the cerebral
hemispheres, concluded that the Australopithecines could speak. Several brain
anatomists, particularly Father Connolly, have challenged his reconstructions.
Moreover, Penfield and his associates have recently discovered that the speech
centers of the human brain lie mostly on inner folds of the cerebral cortex, where
they could not make imprints on the inside of the skull.
Schepers: “The Endocranial Casts. . . .”
Schepers: The Brain Casts of the Recently Discovered Plesianthropus Skulls ,
in Broom, Robinson, and Schepers: “Sterkfontein. . .
Connolly: op. cit.
W. Penfield and L. Roberts: Speech and Brain Mechanisms (Princeton: Prince-
ton University Press; 1959).
260
The Earliest Hominids
The smallest cranial capacity for a fossil man is 775 cc., calcu-
lated for the Trinil Pithecanthropus skull 2.2 This leaves, between
the known ranges of Australopithecus and Homo, a gap of about
200 cc., which Vallois calls a Rubicon. At the moment this stream
is still unbridged.
To summarize, the brain cases of the Australopithecines are hu-
man in form but apelike in size, and in the gross morphology of
the brain, as seen dimly through the surface markings on the inner
surfaces of the skulls, there is nothing to indicate that they, or
other creatures like them, could not have evolved into men. The
ancestors of the genus Homo, whoever they were, must once have
had equally small brains, before they began to grow out of the
Australopithecine range.
The Faces of the Australopithecines
I N 1951 Washburn stated that in the course of human evolution
different parts of the body had evolved at different times and at
different rates.3 First came the erect posture; then the perfection
of the upper limbs; next the brain grew larger; and finally the
teeth grew smaller and the bony structures of the face and jaws
which support the teeth in their work of mastication also became
reduced.
In the evolution of the Australopithecines this schedule may not
have been closely followed. The evidence which we have cited
indicates that the erect posture may not have been fully achieved
at the start, and it certainly did not become more perfect in the
later and larger animals; nor did the brain grow according to
this plan. The teeth, as we shall soon see, grew larger instead of
smaller, and so did their supporting structures. These facts do not
disprove Washburn s thesis, but they cast some doubt on the idea
that the South African Australopithecines were evolving into men.
Although some fossil men may have had as massive faces as the
2 F. Weidenreich: “Giant Early Man from Java and South China,” APAM, Vol
40, No. 1 (1945), p. 97.
3 Washburn: “The New Physical Anthropology,” TNYA, Ser. 2, Vol. 13, No. 7
(l95i), PP- 298-304.
The Faces of the Australopithecines
261
early Australopithecines, nevertheless a whole set of features,
partly concealed by a common air of brutality, separates the Aus-
tralopithecine face from that of fossil and living men. These are in
what I call the mask , the region of the eyes and nose.
Fig. 29 From Proconsul to the Smaller Australopithecines to the Larger
Ones. In this comparison of facial profiles, the skull of Proconsul africanus is repre-
sented twice as large, proportionately, as the other two. The lower part of the
occipital bone of Proconsul has been reconstructed, and the crest of Zinjanthropus
has been left out. In these drawings it may be seen that the brow ridges progress
from none to extremely heavy; the prognathism from extreme to great to medium;
the lateral axis of the eye socket from sloping backwards to straight to sloping
slightly forwards. With an elongation of the face and a reduction in prognathism,
the face has moved upward in front of the brain case. Otherwise, the tooth line
would be inconveniently located for an erect bipedal animal. (Drawing of Pro-
consul after LeGros Clark and Leakey, 1951; Sterkfontein after LeGros Clark,
i960; Zinjanthropus after Leakey, 1959.)
In man the distance between the bony orbits (eye sockets) is
variable; fossil men and the more primitive living races have the
greatest interorbital distances. In the Old World monkeys and
apes this distance is small. Much of the pressure of their jaws is
carried upward by the bony structures on either side of the or-
bits, leaving the eyes-nose triangle a compact island between two
262
The Earliest Hominids
Q
CHIMPANZEE STERKFONTEIN ZINJANTHROPUS
Fig. 30 Variations in the Area of Neck-Muscle Attachment in Apes and
Australopithecines. The chimpanzee represents the apes; Sterkfontein, the
earlier and smaller Australopithecines; and Zinjanthropus, the later and larger ones.
Like the Australopithecus rohustus specimens from South Africa, Zinjanthropus has
wide-flaring temporal crests and a sagittal crest, both indicating an exaggerated
development of the temporal muscles and a coarse, tough diet. But the nuchal
crest, under which the muscles of the back of the neck are attached to the
occipital bone, is set low on the skull in the Australopithecines and high on the
chimpanzee. In Zinjanthropus it is set even lower than in Sterkfontein ( Australo-
pithecus africanus) . This progressions suggests, but taken alone does not prove, a
continuous evolutionary progression among the Australopithecines in the acquisi-
tion of the erect posture. We have no occipital bones from Proconsul with which
to extend this comparison.
columns of stress. In the prosimians and New World monkeys,
both the eyes and the nostrils are set farther apart and the nasal
skeleton takes some of the strain. In the chimpanzee (see Fig. 31)
the maxillary sinuses actually invade the nasal territory, separat-
ing the inner nasal passages from the roof of the mouth by branch
air pockets, and reducing the number of bony struts passing from
the palate to the middle of the forehead from three to one.4 In
man the nasal skeleton takes some of the thrust, and the nose
pushes the orbits farther apart than in most catarrhine monkeys
and all apes. When seen from the side, the human orbit, through
variable, tends to extend to the side and rear much more than in
living Old World monkeys and apes, after the fashion of pro-
simians, platyrrhines, and Proconsul. In these respects man is a
relatively primitive primate.
Furthermore, the nasal skeleton stands out from the plane of
the face, not only in man, but also in lemurs, fossil and living
tarsiers, and most of the arboreal monkeys of both hemispheres,
but not in macaques, baboons, or apes. Their nasal skeletons are
flush with the bones to either side of them, or even depressed. The
same is true of the South African Australopithecines. It has
been said that in all these big-jawed animals the nose would be
4 Broom: “Plesianthropus . . . ,” pp. 86-7.
The Faces of the Australopithecines 263
left standing out like an island were the jaws to be reduced; but
this is not true, because the nasal skeleton is flat even at eye level
where the jaw protrusion makes no difference. No amount of jaw
reduction could give Australopithecus a human nose. In the six-
CHIMPANZEE (ADULT MALE)
STERKFONTEIN
Fig. 31 Sections through the
Nasal Passages of Australopithecus,
Ape, and Man. Section through level
of first upper molar. In the chimpan-
zee the maxillary sinuses lie between
the nasal passages and the palate. In
Australopithecus and man the nasal
passages are directly above the palate
and the maxillary sinuses lie to either
side of the nasal passages only. ( Draw-
ings after Broom and Robinson, 1952. )
HOMO SAPIENS (KAFFIR)
and seven-year-old skulls of Taung and Swartkrans the nasal pro-
file is just as flat as in the adults. Nasally the Australopithecines
were more apelike, and in a sense more evolved, than we are.
In man the external nose is a useful part of the speech appara-
tus: it forms a resonance chamber for the amplification of sound
comparable to the air sac of the gibbon and the enlarged voice box
The Earliest Hominids
264
of the howler. Whether or not the Australopithecines could speak,
and I have seen no conclusive evidence that they could, they seem
to have lacked this special human adjunct.
Telanthropus, however, differs from the others nasally.5 It has a
distinct, though small, nasal spine situated at the front of the na-
sal cavity. In the other specimens the spine is located far inside
and its two lateral segments are divided by the vomer, which is
the lower part of the nasal septum. Telanthropus’s nasal passages
have a distinct floor, forming a single plane set at an angle to the
outer surface of the bone. In the other specimens the floor is
rounded and its junction with the outer plane of the bone gut-
tered and indistinct. In these respects Telanthropus seems to have
been more humanlike than the other Australopithecines.
The Australopithecine Jaws
The Australopithecines needed large jaws. It takes
heavy jaw muscles and big teeth to chew the coarse, uncooked,
and probably mostly uncut food eaten by open-country om-
nivores; witness the large jaws and teeth of the ground-living
baboons. The Australopithecine jaws are just as large and heavy
as they needed to be, and their size relative to that of the brain
case is a purely evolutionary matter without implication of rela-
tionship to other primates.
As in apes, the upper jaw is long and deep, but the palate is en-
tirely unapelike and essentially human in shape. Whereas the ape
palate is long and narrow, with the cheek-tooth rows parallel or
even slightly convergent toward the rear, the Australopithecine
palate is continuously arched, with no apelike gaps (diastemas)
between canines and lateral incisors. The key to the difference be-
tween hominid and pongid palate forms is the relative size and
time of eruption of the upper canines. In juvenile apes, as in a
youthful specimen of Proconsul, the palate is shaped much as in
Australopithecus and Homo. The huge canines of the apes are cut
only when the palate is big enough to accommodate them, and
when the animal needs them — just before or along with the third
5 Robinson: “Telanthropus. . . ,” pp. 445-501.
The Australopithecine Jaws 265
molars. In the hominids the canines are small and erupt early,
without altering the shape of the palate.
Apart from size, the chief difference between Australopithecine
and human palates is that in the former the posterior border of the
palate extends far beyond the level of the third molars, whereas in
man this border is in line with the teeth. In this detail the Austra-
lopithecine palate falls halfway between those of gorilla and man.
It is tempting to interpret this feature to mean that the Australo-
pithecines could not have spoken because their pharynxes had not
yet become open as in man (see page 74), but additional evidence
is needed before such a conclusion can be drawn.
The mandible, which is correspondingly large and heavy, is also
human in shape if compared to the chinless jaws of fossil men. As
CORACOID PROCESS
Fig. 32 The Anatomy of the Mandible: Swartkrans,
female. (Drawing after Dart, 1954.)
in man, the mental foramina are high. No specimen has been
found with a simian shelf to brace it, but even if one turns up,
this feature will not make the creature less hominid. In general the
ascending ramus is extremely long, to accommodate the great
height of the upper jaw and face, and it is set at nearly a right
angle to the occlusal plane in all known jaws except those from
Swartkrans. In the latter the angle is wider because the tem-
poral muscles were attached far back on the brain case. For the
same reason the coracoid process, to the tip and anterior border
of which the temporal tendon is attached, points backward more
than in most human mandibles, particularly those of fossil men.
In all Australopithecine specimens the mandibular fossa of the
266
The Earliest Hominids
temporal bone of the cranium, which is the groove in which the
mandibular condyle moves in chewing, is human in form. Along
with the evidence derived from the teeth, this indicates that the
Australopithecines chewed their food in a rotary movement, as do
primitive men, instead of up and down, like apes and most civi-
lized men. One Swartkrans skull, called female by its discoverer,
had such extensive temporal muscles that they met at the top of
its head in a crest which did not, however, stretch gorilla-fashion
backward to its occipital torus. As even in the gorilla such crests
are confined to males, the identification of this skull as female is
highly dubious. It was probably a male skull.
MAKAPANSGAT
TELANTHROPUS 1
Fig. 33 The Austhalopithecine Mandibles.
The Swartkrans and Kromdraai mandibles are larger and
heavier than those of the earlier and smaller Australopithecines.
This difference does not necessarily imply a change in dietary
habits, because it can be equally well interpreted as an applica-
tion of the law of allometry, that is, that the larger an animal be-
The Teeth of Australopithecus 267
comes, the larger his jaws grow both absolutely and proportion-
ately.
The two Telanthropus mandibles, found in the Swartkrans cave
in what seemed to be a separate pocket of breccia, are more
slender and smaller than those of their companions in the cave,
A. robustus, but they are not identical. The first one is virtually
whole except that both ascending rami are broken off. However,
the position of one condyle was indicated by the form of its
stump, and when reconstructed the ramus height appears to have
been moderate, as in man. Yet the ascending ramus starts to leave
the body of the mandible at the level of the first lower molar, as in
proper Australopithecine jaws, instead of farther back, at the level
of the third molar, as in most human jaws. Also the maxilla that
Robinson attributed to Telanthropus, although manlike in other
respects, was long from nose to tooth line and would require a
high ramus.
The second Telanthropus mandible, which in 1952 Broom and
Robinson tentatively identified as human, is a piece of the right
branch, about 5.5 cm. long, carrying the first and second molars.
Morphologically and metrically it is indistinguishable from a hu-
man jaw fragment and in fact resembles in many ways the Mauer
(Heidelberg) jaw from the Early Middle Pleistocene of Europe.
The Teeth of Australopithecus
More has been written about the teeth of these creatures
than about the sum of the rest of their remains, and much of the
writing is highly detailed and technical, and comprehensible only
to specialists. Luckily the entire subject has been exhaustively
covered by Robinson in a single volume.6
The total number of Australopithecine teeth known in 1956
was 526. Of these 448 are permanent and 78 deciduous, or milk,
teeth. The vast majority come from two sites, Sterkfontein and
Swartkrans, although there are enough from the other sites to in-
dicate what they are like. Every tooth of both upper and lower
jaws, and of both milk and permanent dentitions, is represented
6 Robinson: ‘The Dentition of the Australopithecinae.”
268
The Earliest Hominids
by at least two specimens; and enough specimens of most kinds
of tooth in the permanent dentition are available to permit de-
tailed statistical analysis. These teeth are more abundant and
more completely represented than those of any kind of fossil man
except the Upper Paleolithic folk of Europe, whose teeth were
modern.
TABLE 7
NUMBERS OF
AUSTRA-
LOPITHECINE TEETH
Milk Permanent
Total
Taung
20
4
24
Sterkfontein
12
129
141
Makapansgat
2
25
27
Swartkrans
38
273
311
Kromdraai
6
17
23
78
448
526
1-1 1-2
C
P-1
P-2
M-l
M-2
M-3
Permanent/ uPPer
11 14
22
49
36
49
43
31
(lower
9 5
15
27
25
46
33
33
MiIk/uPPer
4 2
2
5
8
[lower
5 8
9
14
21
Although some of these teeth are very large by human stand-
ards, they are essentially, if not entirely, human in form and func-
tion and quite different from those of either Old World monkeys
or apes, whether fossil or living. These differences are just as
marked in the milk as in the permanent dentitions. In both upper
and lower jaws the incisors are practically identical with those
of Homo both in size and in shape. The canines, too, are similar
in form, but some are larger than the largest found in man. Start-
ing with the first premolar of each jaw, the cheek teeth grow larger
and larger, compaied to mans, as we move down the row to the
third molar, but the morphological similarity remains. No trace is
found in any site of elongated, conical canines, shearing lower
first premolars, or a diastema; nor are any of the molars bilopho-
dont. There is no evidence to indicate a transition from ape to
hominid nor a close relationship with the Old World monkeys. But
the lower molars have the typical Dryopithecus cusp pattern in
common with Proconsul , the other Dryopithecines, and the living
The Teeth of Australopithecus
269
Fig. 34a Australopithecine Teeth: Incisors. A. Sterkfontein 25a; B. Sterk-
fontein 25a; C. Makapansgat (no number); D. Swartkrans 3; E. Swartkrans 68;
F. Swartkrans 11. The smaller and earlier Australopithecines are represented by
only two upper median incisors, a pair in a single jaw from Sterkfontein. One is
depicted here (B), along with an upper lateral incisor accompanying it, and an
upper lateral from Makapansgat (C). An upper median (E) and two upper
laterals (D and F) from Swartkrans represent the larger and later Australo-
pithecines. In A, B, and C a certain amount of surface relief is shown on the inner
or labial side of the blade; this relief takes the form of a mild shoveling ( A and C )
and ridging ( A and B ) . These features also occur in human teeth and will be ex-
plained later. In the Swartkrans median incisor ( E ) , the edges are slightly raised
and the lower border of the tooth is scalloped; this scalloping also occurs in hu-
man teeth when first cut. The Swartkrans laterals ( D and F ) show some ridging,
and each has a teatlike basal protuberance, also found in some human teeth. On
the whole, the earlier incisors seem to have more relief on the lingual side than the
later ones, but the samples are too small to be sure. (Drawings after Robinson,
1956.)
apes. In short, the Australopithecines were just as hominid den-
tally as we are, and in some respects even less apelike.
But they were not a single unit. In tooth size and tooth form
the Australopithecines fall naturally into three groups, just as
270
The Earliest Hominids
their postcranial bones and skulls do, as follows : Australopithecus
africanus (Taung, Sterkfontein, and Makapansgat); Australo-
pithecus rohustus (Swartkrans and Kromdraai); and Telanthro-
pus, known so far only by the lower teeth of two jaws. The Telan-
thropus teeth are indistinguishable from those of Homo.
t> E
Fig. 34b Australopithecine Teeth: Canines. A. Swartkrans 93; B. Sterkfontein
52a; C. Makapansgat (uppers); D. Sterkfontein 50; E. Sterkfontein 51 (lowers).
Like the incisors, the Australopithecine canines show a considerable amount of
relief on the lingual side, including raised edges, ridges, basal protuberances. The
ones shown here are from the smaller and earlier group except A, which is from
Swartkrans. Apparently the smaller and earlier species had more elaborate
relief patterns than the later and larger one. All these variations can be found in
man.
Robinson, who had large enough samples of the first two groups
to permit refined statistical analysis, has shown them to repre-
sent genetically different populations that probably differ as
much from each other as A. africanus does from Homo. On this
basis the two groups should rank at least as separate species.
The Teeth of Australopithecus 271
The differences between them begin with the palate. The pala-
tal index, a length-breadth ratio, falls between 90 and 96 per cent
in the Australopithecines, 63 and 95 per cent in man, and 35 and
62 per cent in apes. These figures place the Australopithecines at
the top of the human range. Although the two species are the
same in this index, they differ in the shape of the upper dental
row and consequently of the palate itself. In Sterkfontein (A.
africanus) the tooth row consists of a smooth curve, as in most
human palates, but in Swartkrans (A. robustus ) the incisors form
a flat line from canine to canine.
In both species the teeth are crowded. Some of the incisors are
set crookedly in the jawbones so that their occlusal edges overlap
each other. All jaws that contain worn incisors have the edge-to-
edge bite typical of human populations that chew tough food,
and the individual teeth are worn not only on their crowns but
also fore and aft, where the teeth rub together in chewing. But in
A. africanus the wear on the lower premolars and molars is mostly
on the buccal or cheek ( outer ) side of the crowns, and in A. ro-
bustus it is on the inner or tongue side, because the palate of
A. africanus is wider than the lower jaw; in A. robustus it is the
opposite. In this feature A. africanus resembles man, and A. ro-
bustus the ungulates. Furthermore, only in A. robustus does the
enamel of the crowns show extensive chipping, as if from grit
encountered on uncleaned roots.
In the length and breadth dimensions of the dental crowns
(see Table 8), A. africanus falls within the human range in fifteen
of thirty-two measurements, overlaps that range in sixteen others,
and falls completely outside it in only one, the breadth of the sec-
ond lower premolar. A. robustus, however, falls outside the human
range in ten measurements, all in the premolars and molars, and
seven of them are breadth measurements. All the ranges of the
two Australopithecine species fail to overlap, but the older spe-
cies, A. africanus, has larger front teeth — incisors and canines —
than the younger species, A. robustus, which has the larger pre-
molars and molars. In this sense A. africanus is more nearly hu-
man than A. robustus. Robinson explains this as follows: “All the
features of the Paranthropus (read Australopithecus robustus )
dentition, as far as size and proportion are concerned, may be ex-
272
The Earliest Hominids
TABLE 8
CROWN DIMENSIONS OF
AUSTRALOPITHECINE TEETH
UPPERS
A. africanus
A. robustus
Zinjanthropus
Garusi
Homo
1-1 1.
b.
9.3- 9.5 a
8.2- 8.3
8.3- 10.8 a
7.3- 7.8
9.6-10.2
7.9
6.5-10.8
6.2- 9.0
1-2 1.
b.
5.8- 7.3 R
5.6- 7.0
6.5- 9.0 «
6.3- 7.6
6.2- 7.1
5.6- 7.3
5.0- 9.0
5.0- 8.5
C 1.
b.
8.8- 9.9 R
8.7- 9.9
8.1-10.6 a
8.4-10.4
7.9 L
9.0-10.2
10.5 ca. s
5.8-11.0
5.0-10.0
P-1 1.
b.
8.5- 9.4 E
10.7-13.9
9.0-10.8 a
13.1-15.3
10.7 L
14.9
9.6 s
12.3
5.5- 9.5
5.0-12.5
P-2 1.
b.
7.2-10.5 E
12.5-13.8
9.2-11.8 a
13.7-16.3
10.7 L
14.9-15.8
9.1 8
12.5
7.8-13.1
5.0-12.5
M-l 1.
b.
11.9-13.2 «
13.2-14.1
13.1- 14.5 a
13.2- 16.6
14.7 L
17.0-18.1
7.8-13.5
9.0-14.8
M-2 1.
b.
12.8-15.1 a
14.3-17.1
13.6-15.9 R
16.0-17.4
17.0 L
18.1-20.3
7.0- 13.6
7.0- 15.2
M-3 1.
b.
11.6- 15.2 »
14.6- 17.9
13.9-17.0 a
15.7-18.1
12.7 L
20.3-21.5
(10.9) 8
(13.0)
4.0- 13.0
4.0- 15.0
L = Leakey
M = Marks
R = Robinson
S = Senyiirek
W = Weidenreich
Nature, Aug. 15, 1959
IJNS, Vol. 109, 1953
TMM, No. 9, 1956
Belleten, Vol. 19, No. 73, 1955
APAM, Vol. 40, Pt. 1, 1945
plained on the assumption that selection has retained as large a
chewing area in the grinding teeth as is consistent with reducing
jaw size at the expense of the less important teeth in a large vege-
tarian.” 7
In one particular comparison the essential difference between
the two Australopithecine species is seen in sharp focus; that is,
the ratio between the crown area of the canine teeth and the
crown areas of the two premolars of each jaw. In A. africanus, as
in various human populations living and extinct, the canines are
nearly or just as large as any premolar of either jaw. In A. robustus
7 Robinson: “Dentition of the Australopithecinae,” pp. 148-9.
The Teeth of Australopithecus
273
LOWERS
Olduvai
Homo Telanthropus Child A. afrioanus A. robustus Meg. jav.
3.5- 6.8
4.9- 7.7
6.5
5.9- 6.0
5.9- 6.3 a
6.1- 8.1
5.2- 5.6 a
5.5- 6.7
4.2- 7.5
5.3- 7.6
6.9
7.0- 7.1
7.3 a
6.8
6.1- 6.7 a
6.7- 7.5
7.0-11.8
5.8-10.4
8.2
8.7-10.0
8.0- 9.2
8.5-10.5 a
9.2-12.1
6.9- 8.5 a
7.3- 9.2
4.5- 9.8
5.7-11.2
8.6 »
10.3
9.9-10.4
9.2
9.2-11.8 a
9.0-11.7
9.2-10.5 »
10.0-12.9
8.0-14.1
8.3-13.2
8.4 »
11.2
9.2- 9.3
9.8-10.1 a
11.6-11.7
10.3-12.5 a
12.0-17.0
8.5-10.5 WM
11.0
8.0-15.0
8.3-13.5
11.9-12.1 a
11.9
13.8-15.0
11.5-12.1
13.0-15.1 a
11.2-13.9
12.7-16.1 a
13.0-15.2
14.0-15.1 WM
13.0
6.3-16.0
8.0-13.7
12.1-13.6 R
12.5-13.1
15.0
11.5
14.3-16.8 a
13.2-15.3
15.0-17.4 a
13.9-16.2
14.5 M
13.0
5.8-15.0
4.0-13.0
13.9-14.1 a
12.3-12.4
13.5-16.7 a
12.7-14.8
15.4-18.5 a
12.9-16.5
15.5 M
13.0
L = Leakey
M = Marks
R = Robinson
S = Senyiirek
W = Weidenreich
Nature, Aug. 15, 1959
IJNS, Vol. 109, 1953
TMM, No. 9, 1956
Belleten, Vol. 19, No. 73, 1955
APAM, Vol. 40, Pt. 1, 1945
TABLE 9
A COMPARISON OF THE CROWN AREAS OF
CANINES AND PREMOLARS IN
AUSTRALOPITHECUS AND HOMO
Upper Jaw
Canine to: First Premolar Second PM
A. robustus
A. africanus
Homo
1.316
1.077
1.021-1.065
1.444
1.107
.980-1.026
Lower Jaw
First Premolar
1.395
1.050
.987-1.025
Second PM
1.546
1.109
1.000-1.078
the premolars are a third to a half again as large as the canines.
This is a considerable difference and taxonomically of signifi-
cance. Had we no other information on the Australopithecine
teeth, we could divide the genus into species on this basis.
274 The Earliest Hominids
Fig. 35 Austbalopithecine Teeth: Premolars and Molars. A. Sterkfontein
uppers; B. Makapansgat lowers; C. Swartkrans uppers; D. Swartkrans lowers. A, C,
and D are relatively new teeth; B is worn enough to obscure the cusp pattern.
Although the molars are more or less the same size in both species, the premolars of
Australopithecus robustus are larger and more molarlike than those of Australo-
pithecus africanus. Also the robustus molars consistently have six cusps; the
africanus molars five or six. ( Drawings after Robinson, 1956. )
Fig. 36 Irregularly Shaped Molars of Australopithecus robustus. A. Upper
Second; B. Upper Third; C. Lower Third. All from Swartkrans. (After Broom and
Robinson, and Robinson, 1956.)
In most dental measurements A. africanus shows more indi-
vidual variation than A. robustus, whose teeth seem to have been
selected for a rigorous, special diet; but in the breadth measure-
ments of some of the premolars and molars the latter’s teeth fall
into two groups, large and larger. Robinson interprets this as evi-
dence of sexual dimorphism, which is not noticeable in the denti-
tion of A. africanus.
PROCONSUL AFRICANUS AUSTRALOPITHECUS AUSTRALOPITHECUS
(STERKFONTEIN) (ZINJANTHROPUS)
HOMO ERECTUS HOMO ERECTUS HOMO SAPIENS
(PITHECANTHROPUS 4) (BROKEN HILL) (AUSTRALIAN ABORIGINE)
Fig. 37 The Upper Canine and First Premolar in Australopithecines Apes
and Men. In Australopithecus the front teeth (canines and incisors) are small in
comparison with the cheek teeth ( premolars and molars ) . In Homo the opposite is
true, and in this respect Homo is intermediate between the Australopithecines and
the apes. In this figure are shown the crown surfaces of six pairs of teeth, the upper
left first premolar ( below ) and the upper left canine ( above ) . The crown patterns
reflect degrees of wear more than morphological differences. What is critical here
is the relative crown sizes of the two teeth in each species. In Proconsul africanus ,
as in living apes, the crown of the upper canine is larger than that of the upper
first premolar. In both the smaller and the larger Australopithecines, represented
here by Sterkfontein and Zinjanthropus, the canine is less than one third as large as
the first premolar. In Homo erectus, represented by Pithecanthropus 4 and the
Broken Hill skull (the earliest and latest of the known 11. erectus skulls), the
canine and first premolar are of roughly equal size, as they are in many modern
jaws, particularly among Mongoloids. The specimen of Homo sapiens shown here,
that of an Australian aborigine, illustrates an extreme degree of size differentiation
for our species the upper canine has only about two thirds the crown area of the
upper first premolar. Because these and other dental differences between Australo-
pithecus and Homo are as old as both genera, they must have arisen longer than
700,000 years ago, the date of the earliest known specimens of either genus, ex-
cept possibly that from Lake Tchad, the taxonomy of which remains to be de-
termined. Of the two, the Australopithecine is the more specialized and the human
the more conservative, at least in the feature illustrated here. ( Drawing 1 from
LeGros Clark and Leakey; drawings 2 and 6 from Clark; number 3 from Leakey;
number 4 from Weidenreich; and number 5 from Pycraft.)
The Earliest Hominids
276
Of the morphological differences that separate the two species,
few if any are great enough to be of taxonomic value. Very few
unworn incisors or canines are available. In the three upper
incisors of A. africanus that we have, the inner or lingual surfaces
are braced with raised rims, ridges, and basal tubercles that give
these teeth added strength and also added grinding surface. In
the three upper incisors of A. robustus these architectural com-
plexities are less pronounced. In the upper canines, three teeth of
each species have rims and ridges, and the two available lower
canines of A. africanus are shaped like mittens, with thumbs pro-
jecting from the lateral side. The second upper premolar of
Fig. 38 The Teeth of Telanthbopus. In the cave at Swartkrans, with the
Australopithecine bones, were found two pieces of mandible and a piece of maxilla
of a smaller and more humanlike hominid, Telanthropus. A represents the right
molars of mandible 1; C, the lower first premolar of mandible 2; and B, a lower
first premolar of Australopithecus robustus, found in the same cave, by way of com-
parison. The molars are humanlike in size and shape, and so is the first lower
premolar. In mandible 2 the socket that once held the lower third molar is conical
and shallow, showing that it held a single-rooted tooth, as in man. As Swartkrans
may be Early Middle Pleistocene in date, it is possible that Telanthropus was an
early form of Homo erectus and not an Australopithecine, but this is not certain.
(Drawings after Robinson, 1953.)
A. africanus has two roots; that of A. robustus three, in all cases —
a total of eighteen teeth for A. robustus and ten for A. africanus.
Some of the molars of A. robustus are not rectangular, as they
usually are among primates, but have odd shapes, including
The Early Hominids from East Africa 277
parallelograms and even triangles. Those of A. africanus conform
more closely to the usual form.
Some of the dental features peculiar to A. africanus, and others
common to both South African species, anticipate subspecific
differences found, over long periods, in the teeth of geographical
races of Homo. In most if not all cases they are typical of either
the Congoid (African Negroid) or Caucasoid branches of man-
kind, and differ from the corresponding features in the teeth of the
Australoid and Mongoloid lines. As human racial differences in
dentition will be discussed in the next chapter, they need not be
MEGANTHROPUS
KROMDRAAI
0 1 2cm
L- 1 — I
Fig. 39 The Australopithecine Features of the Lower First Molar of
Meganthropus. On the left is a diagram of the cusp and groove pattern of the
lower right first molar of the Javanese fossil mandible known as Meganthropus
(see page 298); on the right, the corresponding tooth of an Australopithecine from
Kromdraai, South Africa. The two specimens are virtually identical in size, shape,
system of grooves, and cusp locations. Each has six cusps, a rarity among hominids
except Australopithecus robustus (Swartkrans and Kromdraai), in which the
feature is apparently invariable. Robinson uses the close similarity between these
two teeth, along with other features, including the size and form of the roots, to
substantiate his theory that Meganthropus was an Asiatic Australopithecine.
( Drawings after Robinson, 1953. ) c
given in detail here. But they should be borne in mind, particu-
larly throughout our study of the early hominid remains, includ-
ing teeth, which have recently been exhumed in Tanganyika and
Kenya.
The Early Hominids from East Africa
Early hominid remains have been found in three sites in
East Africa: Olduvai Gorge, Garusi, and Kanam. The only finds
The Earliest Hominids
278
that are completely reliable in a geological sense are those from
Olduvai, and they are also the most numerous and diagnostic.
In Bed I, associated with early Oldowan tools, Leakey found many
hominid remains, including pieces of the skeleton of an eleven-
year-old child of unknown sex, lying 27 feet below the top of the
bed and 78 feet above the basalt at its bottom. This find was made
late in i960. At the time of writing the only available information
comes from two short articles, with photographs.8 1 have also been
given the opportunity, by Matthew Stirling and L. S. B. Leakey
himself, to examine some of the specimens and fresh casts, on
February 25, 1961.
I11 1959 Leakey found the now famous cranium of an adult
male Australopithecine, which he provisionally named Z injan-
thropus boiseii ,9 some five feet above the level of the child’s
skeleton, 22 feet below the top and 83 feet above the bottom of
Bed I. He also found, in i960, more Zinjanthropus specimens, and
the cranium of a Homo erectus in Bed II. In 1913 this same gorge
yielded its first human bones — the complete skeleton of a Capsian
man in or under Bed V — discovered by Dr. Hans Reck. In this
chapter we are concerned only with the material from Bed I.
The Olduvai Child
The Olduvai child’s skeleton consists of a broken mandible,
two more or less complete parietal bones, one wrist bone, and
8 Leakey: “New Finds at Olduvai Gorge,” Nature, Vol. 189, No. 4765 (Feb. 25,
1961), pp. 649-50.
Leakey: “New Links in the Chain of Human Evolution: Three Major New
Discoveries From the Olduvai Gorge, Tanganyika,” ILN, Vol. 238, No. 6344
(March 4, 1961), pp. 346-8.
9 From Balad al-Zanj (or Z inj), Arabic for Land of the Ethiopians, extended
locally to mean East Africa. The surname is that of a benefactor.
Fig. 40 [Facing page] A Section through the Pleistocene Beds at Olduvai
Gorge. C. Olduvai Child; Z. Zinjanthropus; MT. Milk teeth; CH.3. Chellian-3
skull. Capsian denotes the original Oldoway man discovered by Hans Reck; all
the others were discovered by Dr. and Mrs. L. S. B. Leakey. The beds are
numbered I through V. Bed I is Lower Pleistocene but post-Villafranchian. Bed II is
Middle Pleistocene; Beds III and IV, Upper Pleistocene. Bed V is post-Pleistocene,
and some of it has slipped down the face of the gorge to the underlying basalt. In
this section, five different sets of skeletal remains are placed as if they lay directly
over each other. Actually some of them are several miles apart. (Drawing after
Arambourg, 1961.)
72 (236 FT.)
59 (194 FT.)
— CAPSIAN
32 (105 FT.)
O
The Child’s Mandible
281
seven finger bones. Also found were six finger bones, two clavicles
(collarbones), twelve bones of a left foot, and “a few teeth” of
what Leakey calls one or more adult individuals. Before describ-
ing these specimens I should state that the difference of five feet
between the location of the child and his companions and that of
Zinjanthropus may indicate a greater time span than this small
vertical distance would suggest. Although the Oldowan stone
tools extend to the bottom of Bed I, the fauna of the level of the
child s skeleton is said to contain genera and species not seen in
the 22-foot level. The future identification of this new fauna is
critical.
The Child’s Mandible
The mandible was found lying bottom up, and its entire
lower portion had been destroyed. On its left side the mandible
extends to a point one centimeter behind the lower second molar,
Fig. 41 The Olduvai Child’s Man
dible. The mandible of the Olduvai
child, found in Bed I of Olduvai
Gorge, was crushed so that the right
side was bent inward. In this drawing
an attempt has been made to restore
it to its original form. ( Drawing after
Leakey in Nature, i960. )
and on the right it is broken off just in front of the rear border of
the first lower molar. The profile of the chin region is moderately
steep, and continuously curved downward and backward, but on
the inner side the body of the bone sweeps backward almost to a
282
The Earliest Hominids
line connecting the rear borders of the second premolars — farther
back than in the Australopithecines or Proconsul. The forward
flanges of the ascending rami are widely separated from the tooth
row and begin to rise at the level of the forward borders of the
first molars. In this feature the mandible resembles those of Pro-
consul and of all the South African Australopithecines, including
Telanthropus.
Before fossilization the mandible was broken in several places,
all in the canine- and incisor-bearing section. This breakage thrust
the left incisors backward and drew the right side of the mandible
inward. The original shape (see Fig. 41), as tentatively restored,
is that of a long, narrow arch, the two halves of which diverge
slightly toward the rear. In this respect it resembles the mandibles
of Sterkfontein and Makapansgat, Oreopithecus, and the known
Dryopithecines of Africa and Asia, including Proconsul africanus,
all of which retained, in different degrees, a generalized, primi-
tive, essentially V-shaped, primate form.
The Child’s Teeth
All the incisors, canines, and premolars, and both first molars
are present, as well as the left second molar. The first molars and
the incisors are well worn, but the second molar is in mint condi-
tion, having just erupted at the time of death. All the teeth are
within the human range in breadth, but the crowns of the canines,
the second premolars, and the second molar are longer (antero-
posteriorly) than those of any specimen of Homo yet found (see
Table 8, page 273).
The incisors are similar to human teeth in size and form, with no
peculiarities foreshadowing those of any individual human race,
and they also resemble the lower incisors of Proconsul. The ca-
nines, larger than those of the South African Australopithecines,
are pointed and somewhat spatulate, as in many human beings.
The premolars are longer than wide, especially the second pre-
molars. Instead of being superhominid, like those of the Australo-
pithecines, the premolars are, if anything, less completely bicus-
The Child's Parietal Bones
283
pid than our own, and each of the second premolars has a pit
(fovea) at the rear of the crown. This pit is bordered posteriorly
by a rim composed of tiny cusps, or beads, as in the corresponding
teeth of Proconsid. The cusp pattern of the first molars is fully hu-
man, with five crowns on the left tooth and six on the right one.
The left second molar has six cusps. Seen from the side, these
teeth fail to form a straight row at the line of occlusion with the
upper teeth: the crowns of the premolars are lower than those of
either the molars or the canines and incisors.
These are clearly not proper Australopithecine teeth. Rather,
they resemble those of Homo in form and also in relative size
along the tooth row; the incisors and canines are large compared
to the Australopithecine dentition, and the molars are relatively
long and narrow. In fact, the molars also resemble, in this respect,
those of Telanthropus, Proconsul, and Oreopithecus. Morpho-
logically the molars look like Proconsul’s rather than Oreopithe-
cus's; the opposite is true of the premolars. Whatever the kinship
and status of the Olduvai child turn out to be, his teeth seem to
form a connecting link between the large apes of the Miocene and
Pliocene and Homo. The living apes and the South African Aus-
tralopithecines would then be left on either side of him. That he
could have been descended from the Australopithecines so far
found seems unlikely, but some of them could have been de-
scended from him.
The Child’s Parietal Bones
With the mandible were found two parietal bones, the left of
which is the more nearly complete. Their open sutures and thin
walls confirm the eleven-year-old age ascribed to the mandible.
According to the scale on the photographs, these parietals are
about as long, in the sagittal chord ( from bregma to lambda ) as
those of Sterkfontein.1 All else being equal, this comparison sug-
gests a capacity of 450 to 500 cc., but this is a very tentative
1 Roughly, 70 to 75 mm. In Homo erectus the shortest chord known is that of
Pithecanthropus 1, 87.5 mm., capacity goo cc. Pithecanthropus 2, with a chord of
91 mm., had a capacity of 775 cc.
284
The Earliest Hominids
figure, one which does not allow for the curvature of the bones. In
1962 Leakey showed a picture of himself placing the parietals of
the Olduvai child over the brain cast of Pithecanthropus 2, which
had a cranial capacity of 775 cc. The bones were almost large
enough to fit. The child’s cranial capacity may, therefore, have
been as high as 700 cc.
The cranial capacity of an eleven-year-old modern child is
virtually that of an adult, and the parietals have assumed nearly
adult form. These parietals are thin, and show no signs of an
incipient crest seen on the larger Australopithecines. The left
parietal contains a depressed fracture with radiating cracks, in-
flicted pre or ad mortem.
The Foot Accompanying the Child’s Remains
The left foot found with the Olduvai child’s mandible
and parietals was at first attributed to the same individual, but
in 1962 Leakey said that it belonged to an adult. Although it is
large enough for a modern eleven-year-old, it is too small for a
living adult above the size of a Pygmy, but it might have been
large enough for Australopithecus africanus.
From a photograph in which they were roughly assembled for
the record only, I have drawn a tentative reconstruction of this
foot (Fig. 25). It is shown in comparison to the foot bones of a
modern Bushman and of a male gorilla, minus the toe bones,
which are missing in the Olduvai specimen. Regardless of the
accuracy of detail, several differences are evident at once, The
Olduvai foot is small but large enough for an eleven-year-old
child. It is shorter and broader in the tarsal region than the mod-
ern foot, and narrower than the gorilla’s. The trochlea (upper
articular facet ) of the talus curves outward in man and in Olduvai
(if this reconstruction is correct) and inward in the gorilla. In
man the navicular is relatively thick anteroposteriorly, and nearly
straight from side to side. In the gorilla this bone is narrow and
curved, with a spur projecting on the inside of the foot. In these
respects the Olduvai navicular is intermediate between man and
gorilla. In the shapes and disposition of the cuboid and cunei-
The Collarbone, Hand, and Fingers 285
forms, the Olduvai foot is also intermediate between man and go-
rilla, although the position of the medial cuneiform is question-
able.
The Olduvai metatarsals suggest a foot fully or almost fully
adapted for walking rather than for grasping, but the axis of the
calcaneus was apparently curved, as in the gorilla and Procon-
sul nyanzae, so that the heel part of the bone seems to be set to
the left of the center line of the foot instead of more directly un-
der the talus, as in man. The outer side of the foot, from meta-
carpals to talus, seems to have borne more of the body’s weight
than in living man, but not as much as in the gorilla. Human feet
are variable in this respect, however, and in others.
The Olduvai talus appears similar in general to Kromdraai’s,
and it and what is left of its calcaneus may, on further study, turn
out to be intermediate in many respects between those of Pro-
consul and man. From the functional point of view the Olduvai
foot was large enough to support the whole body of a child or a
very small man, and so shaped that he could at least have begun
to walk erect. Whether or not he did so is another question.
The Collarbone, Hand, and Fingers
Leakey’s preliminary notices contain pictures of one clavicle,
originally attributed to the child and later called adult. That of
the other adult is not shown. The clavicle shown has lost both
ends, and in its present condition is 13 cm. long, but was proba-
bly at least 2 cm. longer, long enough for a full-sized modern man.
Its shape is a simple, open S, as in man. In the gorilla the bone is
shaped like a hockey stick, with a curve at only one end.
The finger bones are said to come from the same two individu-
als, and there is only one hand bone, a capitate — the most cen-
trally located of the wrist bones. Luckily we have a comparable
bone from Sterkfontein. Seen from the volar or palm side, the
capitate is about 25 mm. long and 20 mm. wide, well within the
human range. Although details are not easily distinguishable, it
seems to lack the constrictions on both sides characteristic of this
bone in Proconsul and the living apes. These constrictions serve as
286
The Earliest Hominids
anchors for the powerful tendons needed to keep the wrist firm for
brachiation. The Sterkfontein capitate also lacks them.
The finger bones of the first individual consist of five first or
second phalanges, all broken at the proximal (wristward) end,
and two terminal or nail-bearing phalanges. The finger bones of
the second, so-called adult individual, are two proximal or inter-
mediate phalanges ( they are not easy to tell apart when broken )
and two distals ( the ones that carry the fingernails ) . Two distals
and one intermediate are intact. The phalanges of the two indi-
viduals are alike in size and shape, and as large as those of some
living men.
They are not, however, fully human in form. The shafts of the
proximal and intermediate bones are broad, flattened, and lipped
on the palm side, as in Proconsul and the gorilla, and their ter-
minal joints are narrower than their shafts, which have convex
borders. In man the joints are usually wider than their shafts, the
borders of which are slightly concave. In these details the Olduvai
finger bones are less human than those of Kromdraai.
But in another respect the distal phalanges of both Olduvai
individuals are human rather than apelike. (We have no distal
phalanges from Kromdraai for comparison.) In man the end of
each distal phalanx is broad and rounded, to support pressure
from a broad, flattish nail, whereas in the apes the distal phalan-
ges are tapering and pointed, to match the narrow, curved nails
that cover them. In this anatomical detail both Olduvai hands
were nearly if not entirely human.
Luckily one piece of thumb bone was recovered. It is the outer
(distal) half of the proximal phalanx of the left hand. Its articular
joint, on which the distal phalanx moves, is as wide as any corre-
sponding joint on the fingers of the same hand, or very nearly so.
This is the human condition. In apes it is narrower than the finger
joints.
In brief, these Olduvai hominids had wrist bones like those of
Kromdraai and man, proximal and intermediate finger phalanges
like those of Proconsul and the gorilla, terminal finger bones like
man’s, and proximal thumb bones of human size compared to the
sizes of the finger bones. The total picture is one of evolution in
process.
The Evolutionary and Taxonomic Position
287
The Evolutionary and Taxonomic Position of
the Olduvai Child
Until the remains of the Olduvai child and his adult com-
panions have been carefully and competently studied by special-
ists in primate and human anatomy, we shall not know where on
the hominid family tree this child belongs, nor what we should
call it. Leakey has shown admirable forbearance in declining to
give it a hastily coined Latin name.
These bones and teeth may be compared to those of Proconsul,
the Fort Ternan primate, the Australopithecines, the living apes,
and men. In many features the child resembles Proconsul, so
much so that, if future studies support my tentative interpretation
of the pictures and hasty handling of the specimens, a case can be
made for the child’s probable descent from a Dryopithecine,
perhaps Proconsul himself. The Fort Ternan primate may be even
closer. The child resembles the living apes only in features which
both share with Proconsul. As in many ways the child is like South
African Australopithecines, he probably belongs to the genus
Australopithecus, but he is at the same time both more dryopithe-
cine and more human than any Australopithecine yet found. He
resembles man enough, perhaps, to have been our ancestor — pro-
vided that Leakey does not unearth some part of him which con-
tradicts this interpretation.
Even more important than finding the rest of the child’s bones
and teeth, however, is determining their geological age. Is there a
soil change in the five feet that separate Zinjanthropus’s lair and
the child’s? What, if any, is the difference in fauna between these
levels? The answers to these questions will help us find out.
Z injanthropus: His Tools, Diet, and Activities
The specimens known collectively as Zinjanthropus ( I am
using the name informally, like Telanthropus) include the 1959
cranium 2 and the i960 discoveries — a tibia, a fibula, parts of a
2 “The Astonishing Discovery of ‘Nutcracker Man’; Dr. and Mrs. Leakey at
Work at Olduvai,” ILN, Vol. 235, No. 6267 ( 1959), pp. 217—19.
Leakey: “The Newly Discovered Skull from Olduvai: First Photographs of the
Complete Skull,” ILN, Vol. 235, No. 6268 (1959), pp. 288-9.
Aranibourg: “L’Hominien Fossile d’Oldoway,” pp. 223-8.
288
The Earliest Hominids
second skull, and some loose teeth. No systematic study of any of
them has yet been published, and of the i960 discoveries only the
tibia and fibula have been illustrated.3
The site was apparently a camping place along the shore of a
lake, at the head of a small peninsula. In it animal bones are abun-
dant, particularly those of snakes, lizards, and crocodiles. Birds
too are common, including a giant ostrich that laid giant eggs.
Some of the mammals are also giants: Afrochoerus was a pig the
size of a rhinoceros; Pelorovis a sheep six feet at the shoulder,
with a horn spread of twelve to fifteen feet; Sivatherium a short-
necked giraffe with horns like moose antlers; and Simopithecus a
baboon with a lower jaw the size of a gorilla’s.4
The hominids whose bones were found in the 22-foot level of
Bed I seem not to have been skilled hunters, if they were hunters
at all. Zinjanthropus 1, who died just after erupting his third mo-
lars, had already worn some of his other grinding teeth down to
their pulp cavities. To have achieved this degree of attrition he
must have been eating gritty roots, extracted from the soil, per-
haps with sticks sharpened by his stone tools, and he may have
eaten them uncleaned.
Most of the reptiles and some of the small mammals fall into the
category of “slow game,” game usually killed by women and chil-
dren among living food-gatherers. The representatives of the giant
mammals were apparently sucklings or even newly born babies,
which are also sometimes killed and eaten by baboons. Leakey’s
careful study of these bones indicates that Zinjanthropus could
not have been the hunter that Homo was, as shown by every pre-
agricultural living site attributed to the latter which has yet been
found. Yet the Zinjanthropi indubitably ate more animal proteins
than were consumed by any known ape. They must have already
developed a taste for raw meat.
3 Leakey: “Recent Discoveries at Olduvai Gorge,” Nature, Vol. 188, No. 4755
(i960), pp. 1050-2.
Leakey: “New Links in the Chain of Human Evolution: Three Major New
Discoveries from the Olduvai Gorge, Tanganyika,” ILN, Vol. 238, No. 6344
(1961), pp. 346-8.
The Zinjanthropus cranium has been studied by P. V. Tobias, whose volumi-
nous report is expected shortly.
4 Leakey: “Finding the World’s Earliest Men,” NG, Vol. 118, No. 3 (i960),
pp. 420-35.
The Anatomy of Zinjanthropus: His Cranium 289
Zinjanthropus was an Australopithecine. So were the hominids
who lived in South Africa at roughly the same time. If he could
not hunt full-sized, adult game, neither could they. This new evi-
dence effectively lays the ghost of Australopithecus the Hunter
conjured up by the juxtaposition, in the Transvaal breccias, of
hominid and other mammalian bones, including brained ba-
boons.
The Anatomy of Zinjanthropus: His Cranium
The cranium of Zinjanthropus 1 has been pieced together
and restored enough so that we can see what it was like. Its most
striking feature is an exuberant growth of bony struts to brace the
movements of a pair of massive jaws. The face is enormously long,
and the brow ridges rise above the level of the forehead, which
slopes backward, at first, behind them. On the sagittal line of the
brain case rises a crest, as in Swartkrans, to anchor the temporal
muscles where they meet on top of the skull. This crest is split
down the middle by the sagittal suture, which had not yet fused at
the time of death. Having just cut his wisdom teeth, Zinjanthropus
was barely eighteen, according to our human growth schedule,
and as an Australopithecine he may have been even younger.
Other crests run backward across his temporals from the zygo-
matic arches, and below them is set a pair of man-sized mastoids.
In the rear his neck muscles were accommodated by still another
crest, which is set low down, indicating an upright hafting of the
skull on the neck.
Zinjanthropus s face, although larger and more apelike in gross
proportions than the faces of the other Australopithecines, is more
human in three respects. His eye sockets are wide and set far
apart, so that their lateral borders are cut back, as in both Procon-
sul and man. His nasal skeleton can be seen from the side along its
entire length, whereas in the other crania it is recessed. The nasal
spine is set forward, at the lip of the nasal floor, as it is in Telan-
thropus alone of the South African Australopithecines.
Leakey has published a few of the measurements of this cra-
nium. I have tentatively added others based on all the available
photographs, and I have compared them with figures similarly ob-
290
The Earliest Hominids
tained for Australopithecus africanus, A. robustus, and Proconsul
africanus (see Table 10). Let us first consider only the three
Australopithecines. In twelve of fifteen measurements, a progres-
sion from smallest to greatest may be seen from A. africanus to
A. robustus to Zinjanthropus. In two other measurements no fig-
ures are available for A. robustus, but the progression is otherwise
valid. In only one measurement, head height, is the sequence re-
versed; Sterkfontein has the highest cranial vault and Zinjanthro-
pus the lowest.
In these changes — assuming that we have an evolutionary se-
quence— the cranial vault is affected the least, and most of the
growth is seen in the bony framework supporting the jaws. The
greatest increase is evident in the dimensions of the face; upper
face height, bizygomatic face breadth, nose height and breadth,
orbital height, and palate length and breadth.
On the other hand, changes in proportions, as reflected in seven
indices, are trivial or nonexistent in four, and probably significant
in only three — the two height ratios of the vault and the orbital
index. These simply mirror the flattening out of the vault, the
great growth of the jaws, and the change from more apelike to
more human facial proportions. The eyes are farther apart than in
the other Australopithecines and apes, and the stress of jaw action
passes more through the center of the face and less through its
sides than in the South African crania. The reduction in facial
flatness seen in Zinjanthropus not only harks back in a sense to
Proconsul but also makes him look more human.
As for Proconsul, we can see on Table 10 that P. africanus ,5 the
smallest ape of this genus, is much smaller in all dimensions than
the Australopithecines but is similar in several cranial and facial
dimensions. Its high length-breadth (cranial) index is due pri-
marily to its lack of brow ridges, which stretch the cranial lengths
in the Australopithecines. Its facial proportions, involving the up-
per face, nasal bones and apeture, orbits, and palate, are very
similar. In the assessment of genetic continuity in human geo-
graphical lines these ratios seem to be more important than gross
5 We are obliged to limit our comparison to that species because no skulls of
the other two have been found, only jaws and teeth.
The Anatomy of Zinjanthropus: His Cranium
291
TABLE 10
TENTATIVE CRANIAL MEASUREMENTS
AND INDICES OF THE AUSTRALO-
PITHECINES AND OF PROCONSUL
AFRICANUS
Aa to Z
Proc. af. A. afric. A. rob. Zinj. Directions
Maximum Length (glabello-
occipital)
97??
147 H
174 L
HA
Maximum Breadth (supra-
mastoidal)
120
h-)
GO
CO
Maximum Breadth (inter-
temporal)
83
99 H
116
118 L
>
Basion-bregma Height
Aur.55 +
105 H
102
99 L
<
Bizygomatic Face Breadth
87
131
137/154 «>
188 L
»
Biorbital Diameter
70
88
102
122
>
Interorbital Diameter
16
24
39
42
>
Upper Face Height (nasion-
prosthion)
45
74
86
114
»
Nose Height
24
49
61
73
>
Nose Breadth
14
27
32
42
A
Maxillary Height (nasale-
prosthion)
26
28/30 «>
37
>
Orbital Height
21
33
34
37
>
Orbital Breath
27
33
37
46
y
Palate Length
50
64.6 <»
70
84
— >
Palate Breadth
50?
64.6 «>
68
82
— ♦
Cranial Index
86?
67
68
0
Length-Height Index
58?
71
57
<-H
Breadth-Height Index
67
106
87
84
<
Upper Facial Index
53
56
63/56 O)
61
O
Nasal Index
56
55
52
58
O
Orbital Index
78
100
95
80
<
Palatal Index
100
100
97
98
O
H = G. Heberer: Primatologia, Vol. I (1956), pp. 379-560.
L = L. S. B. Leakey: Nature, December 17, 1960.
(2) = two specimens (5) = mean of 5 skulls: 4 Sterkfontein and 1 Makapansgat
dimensions, and the same may be true of the African primates in
question.
Returning to Zinjanthropus, we note that in a preliminary esti-
mate Leakey has set the cranial capacity of that skull at over
600 cc. If he is right, it might fall within the maximum pongid
figure of 685 cc. for a male gorilla. If we postulate a generous
maximum of 700 cc. and calculate a ratio between brain size and
292
The Earliest Hominids
palate area, the figure for Zinjanthropus appears to be between
1.1 and l.o, compared to 1.2 for a female chimpanzee. Homo
erectus ranges from 1.4 to 1.7, and Homo sapiens from 1.9 upward.
If we study the outlines of the crania of the smallest species of
Proconsul, P. africanus, and of Sterkfontein and Zinjanthropus,
as shown on Fig. 29, and consider them, at least for the moment,
an evolutionary sequence, the metrical progression indicated on
Table 10 takes on added meaning. The position of the skull in re-
lation to the neck shifts from the diagonal hafting of a pronograde
ape to the vertical one associated with the erect posture. The face
becomes more vertical also, as it moves higher and higher up on
the front of the brain case : if the face grows more rapidly than the
brain it has to expand upward as well as downward, resulting in a
loss of forehead. Compared to the Miocene ape Proconsul, the
Lower Pleistocene Australopithecines have grown progressively
larger-faced and more brutal-looking. Had we a sequence of the
pongid line that led from Proconsul to gorilla, a parallel progres-
sion could presumably be seen.
This exercise does not mean that, if our premise is true, Zin-
janthropus was evolving away from a common ancestor with man.
It simply shows that man’s face had to grow larger before it could
become small again. This up-and-down sequence could reflect
merely an alternation of increasing chewing needs, which began
with a dietary shift from fruit to roots and raw meat, followed by
a decrease brought about by the invention of cooking. We shall
see a repetition of this rise and fall of the facial scaffolding in
several human sequences. Here the important thing is to estab-
lish the principle that governs this kind of change.
The Teeth of Zinjanthropus
The only teeth from the 22-foot level of Bed I of Olduvai
Gorge about which anything has been published are the sixteen
6 Sir Arthur Keith, who invented this index, compared brain size to palate area
irectly. I have treated brain size as a cube and palate area as a square, dividing
the cube root of cranial capacity by the square root of palate area, and thus have
compiled new figures for all fossil skulls found since he wrote. A. Keith: The
Antiquity of Man, Second Edition (London: Williams and Norgate; 1925), Vol I
pp. 213-6.
The Teeth of Zinjanthropus 293
uppers in the 1959 skull. We have all sixteen. The third molars had
been cut but had not yet descended to the occlusal plane and so
were not worn at the time of death. These teeth are similar to
those of the South African Australopithecines in size and shape,
with a few exceptions (see Table 8). The incisors fall within the
South African range, and the canines are even smaller; the left
canine is reduced to a small, degenerate-looking cone. The pre-
molars match those from the south, but the molars are larger, the
second and third having particularly excessive breadths.
With these molar breadths we can arrange a progression from
A. africanus to Zinjanthropus comparable to that for the cranial
measurements, with the same implication. Species of a single
genus grow progressively larger-toothed and heavier-jawed as
they grow bulkier, and even more so as they meet increasing
needs for processing coarser and tougher items of diet. A similar
progression can be seen in the Proconsul series, among the Indian
god-apes, and indeed in many other kinds of mammals. Among
the pongids and hominids this increase in tooth size tends to in-
volve a widening rather than an elongation of the cheek teeth, be-
cause in a semi-erect or erect animal there is more room in the
jaw for lateral than for longitudinal expansion, and the more
erect an animal stands, the more cumbersome to him a long muz-
zle becomes.
Morphologically the upper teeth of Zinjanthropus are Austra-
lopithecine, in the South African sense, with a few differences.
Although even larger than those of A. rohustus, the Zinjanthropus
molars are generally rectangular rather than irregular in shape,
recalling those of A. africanus in this sense. The barely erupted
third molars are relatively short-crowned, and their enamel is
extensively and finely wrinkled, as in the molars of the orang.
Compared with the teeth of the Lower Miocene apes of Kenya,
Zinjanthropus’s molars recall those of Sivapithecus africanus
rather than those of Proconsul, for the crown patterns of the
teeth of S. africanus are relatively simple. Compared with the
teeth of Homo, Zinjanthropus’s exceed the human range in
breadth from canines to third molars, and in the length of the
first premolar and the first and second molars. In size and in most
294
The Earliest Hominids
proportions Zinjanthropus s teeth are the least human of the Aus-
tralopithecine teeth so far discovered.
The Leg Bones Attributed to Zinjanthropus
A TIBIA and a fibula, presumably a pair, were found in the
22-foot level of Bed I in Olduvai Gorge, where the Zinjanthropus
skull lay. These bones are long, very slender, apparently straight,
and broken at the lower ends. As far as one can tell from a single
photograph, they appear to be essentially human, but the fibula
is relatively heavier than the tibia. They must come from two
individuals, or else an unusual amount of weight was carried on
the outer margin of the foot.
As more than one skull has already been found in this level,
there is no reason for attributing these leg bones to the first
Zinjanthropus skull, and indeed they appear too slender to go
with it. If these were the bones of a human being, he, or more
likely she, would have had a stature of only about 136 cm., or four
feet six inches, more or less. If they were the bones of a Zinjan-
thropus whose legs might have been short in proportion to trunk
length, a stature of five feet is possible. However, until further
studies are made, we can do no more than speculate about the
significance of these leg bones.
The Status of Zinjanthropus
We may provisionally conclude that Zinjanthropus was an
Australopithecine, more manlike in some respects than the Aus-
tralopithecines of South Africa and less so in others; that in the
development of his teeth and of the supporting bony structures
of his face and brain case he came at the peak of a divergent
evolutionary line; and that his relationship to the earlier Olduvai
child, who so far is represented only by bones and teeth missing
in Zinjanthropus, was not so close as a stratigraphic distance of
only five feet would imply. These two denizens of Olduvai seem
to stand at opposite ends of the Australopithecine scale, if the
child is, in fact, an Australopithecine.
The Kanam Mandible
295
The Specimen from Lake Eyasi, Tanganyika
I N J93^> when the study of the Australopithecines was in its
infancy, L. Kohl-Larsen, a German paleontologist, found another
specimen at Garusi, on the eastern shore of Lake Eyasi in Tan-
ganyika, about 35 miles south of Olduvai Gorge. It lay in a Lae-
tolil faunal bed, where it might possibly have been intrusive. It
consists of a small piece of left maxilla containing two premolar
teeth. A third molar picked up a few miles away and originally
attributed to this specimen or one like it was probably human and
less ancient.
Kohl-Larsen originally called it Australopithecus, with a ques-
tion mark; in 1950 Weinert dubbed it Meganthropus africanus
( the name Meganthropus had been previously held by a Javanese
specimen that will be described shortly); and in 1955 §enyiirek
labeled it Praeanthropus africanus. Also in 1955, Robinson
showed, to the satisfaction of most primate paleontologists, not
only that Garusi is an Australopithecine but that it is indistin-
guishable from Sterkfontein, with which it may have been con-
temporary. ‘ In any case, subsequent discoveries have rendered
the age and taxonomy of the Garusi specimen unimportant.
The Kanam Mandible
The same may almost be said of the famed Kanam mandibu-
lar fragment discovered in 1932 by a Kikuyu assistant of Leakey.
It lay in an Omo faunal deposit on the south shore of the Kavi-
7 L. Kohl-Larsen: Auf den Spuren des Vormemchen, Vol. 2 (Stuttgart: Strecker
and Schroder; 1943), pp. 379-81.
H. Weinert: t)ber die Neuen Vor-und Friihmenschenfunde aus Afrika, Java,
China, und Frankreieh,” ZFMUA, Vol. 42 (1950), p. 113-48.
A. Remane: “Die Ziihne des Meganthropus africanus,” ZFMUA, Vol. 42
(1951), PP- 311-29-
Robinson: Further Remarks on the Relationship between *Meganthropus> and
Australopithecus africanus,” AJPA, Vol. 13, No. 3 (1955), pp. 429-45.
M. S. Senyiirek: A Note on the Teeth of Meganthropus africanus Weinert
from Tanganyika Territory,” Belleten, Vol. 19, No. 73 (1955), pp. 1-55.
Heberer: “Die Fossilgeschichte . . . ,” pp. 379-560.
The Earliest Hominids
296
rondo Gulf of Lake Victoria Nyanza, in Kenya, about 200 miles
north and a little west of Olduvai Gorge. It lay in what appeared
to be basal Villafranchian soil, associated with pebble tools and
a tooth of the extinct Dinotherium .8 For nearly thirty years this
fossil has been the center of controversy, regarding both its age
and the kind of hominid it was.
Its age was questioned because the site eroded away after the
specimen was removed, and could not be relocated. Later, when
fluorine and uranium tests were applied to the specimen, it was
found to have a high calcium carbonate content, which invali-
dated the comparisons made between it and other specimens of
the same period and region. It is very old, but its exact date is un-
known.9
The other source of doubt was its morphology, because it
seemed to have small teeth and a chin. The fragment consists of a
battered and diseased piece of lower jaw extending from the
distal root of the right first molar to the region of the left second
premolar. Only the two right premolars, both badly worn and
broken, are in situ. Also, the lower margin of the mandible is
missing. After an extensive study of this amorphous-looking speci-
men, Tobias found that it had no chin at all, that the protuberance
resembling one was a bone sarcoma that had grown over an old
fracture. Its greatest distinction is the massiveness of the bone,
which exceeds those of all known jaws of Homo in at least one
dimension, symphyseal height.1
Kanam man was either an Australopithecine contemporaneous
with the other animals found with him; or he was a later Aus-
tralopithecine intiusive in the deposit from which he was removed;
or he was an equally intrusive Homo comparable in age and
grade to those found in North Africa, which will be described in
Chapter 12. As we now have a human cranium from the Early
8 Leakey: The Stone Age Races of Kenya (Oxford: Oxford University Press-
1935)-
9 Oakley: Phtjsical Anthropology in the British Museum, 1958; and personal
communications .
1 P' V- Tobias: “The Kanam Jaw,” Nature, Vol. 185, No. 4714 (i960) pp
946-7- ’
Tobias’s definitive report on the Kanam mandible will be published in 1962,
in the Transactions of the Fourth Panafrican Congress on Prehistory, held in
Leopoldville in 1959.
The Fossil Hominid of Tell Ubeidiya 297
Middle Pleistocene of Olduvai Gorge, it makes little difference to
the history of man and the Australopithecines which of the three
he was.
The Australopithecine from the Republic of Tchad
On June 3, 1961, Yves Coppen, a French paleontologist, an-
nounced his discovery of an Australopithecine skull in an un-
named Lower Villafranchian site in the Republic of Tchad, half-
way between Largeau and the Nigerian frontier and about 200
miles northeast of Lake Tchad.2 The specimen is called Lower
Villafranchian because of its association with an extinct elephant,
Loxodonta africanavus, so named by its discoverer. If this dating
is substantiated after a complete study of the fauna, the Tchad
Australopithecine may turn out to be the oldest of its genus.
The skull fragment consists of a frontal bone broken off a short
distance in front of bregma, and parts of the bones of both sides
of the face, the right of which is the better preserved. The skull has
a forehead, and its cranial capacity was apparently large for its
genus. The brow ridges are of moderate dimensions, and overlay
large frontal sinuses. The orbits are very large, and over each of
them is a supraorbital foramen, a human feature. The zygomatic
bone (malar) is short and thick; the junction with the zygomatic
arch is sharply curved and sloped obliquely upward and forward.
The lower face is extremely prognathous, but there is a canine
fossa, and the canine teeth were apparently small. Coppen provi-
sionally considers this to be the oldest and the most nearly hu-
man of the Australopithecine specimens yet found.
11
The Fossil Hominid of Tell Ubeidiya, Jordan Valley
The fossil hominid found in a Lower Pleistocene outcrop at
Tell Ubeidiya, Israel, near the southern shore of Lake Tiberias
and on the west side of the Jordan Valley, consists of two small
2 Y. Coppen: “Decouverte d’un Australopithecine dans le Villafranchien du
Tchad,” CRAS, Vol. 252, No. 24 (1961), pp. 3851-2.
298
The Earliest Hominids
pieces of skull and one incisor tooth. We do not yet know whether
this animal was Australopithecus or Homo. Although the speci-
mens await description, the skull fragments are said to be “of
very great thickness,” 3 and from other sources I have heard that
the incisor is small.
As stated earlier, the tools also found are similar to those from
the Zinjanthropus level at Olduvai Gorge and from Ain Hanech
in Algeria.
The animal bones are those of fish, turtles (including terra-
pins and tortoises), birds, and mammals. Many slow-moving and
verminous animals, such as tortoises, mice, other rodents, and
porcupines, were apparently eaten, but so were large mammals.
Hippopotamus, rhinoceros, elephant, asses, zebras, a large cervid
deer, a fallow deer, and gazelles and antelopes were all, it seems,
on the menu, however their flesh was acquired. Some of the larger
animal bones were split, presumably for marrow, and one of them
showed scratches where flint may have been used to cut off the
flesh. Still, most of the bones were those of slow game. If the fossil
hominid of Tell Ubeidiya was a better hunter than Zinjanthropus,
his superiority in this respect has yet to be definitely proved.
The fauna is a combination of Oriental, Palearctic, and African
genera. The two kinds of deer, one of which, the fallow deer
( Dama cf. mesopotamica) , still exists in the Near East, are Pale-
arctic whereas the zebras are African. The hippopotamus is both
African and Oriental; one pig ( Sus cf. scrofa ) is Palearctic, an-
other has not yet been identified. A fresh-water turtle ( Trionyx )
has living species in both Africa and Asia. Because Palestine
stands at the crossroads of continents and faunal regions in the
Old World, it could have served as a link between the Aus-
tralopithecines of Africa and those of Asia, wherever they origi-
nated and to whichever regions they subsequently dispersed.
The Meganthropus Mandibles from Java
That the Australopithecines dispersed widely is evident
from the discovery of two mandibular fragments in the Djetis
3 Stekelis et al. : “Villafranchian Deposits Near Ubeidiya . . . p. 182.
The Meganthropus Mandibles from Java 299
faunal beds of Java, known as Meganthropus paleojavanicus. Von
Koenigswald found the first in 1941 and P. Marks, a Dutch
geologist, retrieved the other in 1952. Both came from Sangiran,
the site of the infant human skull that von Koenigswald calls
Pithecanthropus modjokertensis ; and at least the first one came
from the same level as the human specimen.4 More clearly than in
South Africa, this evidence indicates that two kinds of hominid
were sympatric, if only for a short period.
The first piece (see Fig. 56, p. 381) includes the first lower
molar, both premolars, the socket of a canine, and a small section
of the inner sagittal surface. The second contains the premolars
and molars of the right side of the mandible; but the crowns are
broken off or abraded, except that of the third molar, which is
intact.
Both jawbones are large and thick, well outside the human
range but within that of the larger South African Australopithe-
cines. Morphologically the Javanese and African mandibles are
similar but not identical.5 The teeth, which have been widely dis-
cussed, fall within the South African size ranges in length and
breadth, and all but the lower first premolar are closer to Aus-
tralopithecus africanus than to A. robustus ( see Table 8 ) .
In seven of ten dimensions, these teeth are also within the
ranges of Homo. The only two molars that have crowns, the first
lower molar of von Koenigswald’s specimen and the third of
Marks’s, are also closest to A. africanus in shape. Von Koenigswald
has advanced several arguments to show that the teeth of his
specimen are more nearly human than those from South Africa,
but this can hardly apply to Telanthropus or to the newly dis-
4 Weidenreich: op. cit.
P. Marks: “Preliminary Note on the Discovery of a New Jaw of Meganthropus,”
IJNS, Vol. 109, Nos. 1, 2, 3 (1953), pp. 26-33.
Robinson: “Further Remarks on the Relationship between ‘Meganthropus’ and
Australopithecines,” AJPA, Vol. 13, No. 3 (1955), pp. 429-46.
Von Koenigswald: Meeting Prehistoric Man (New York: Harper & Bros.;
1956).
5 Von Koenigswald finds, on the inner side of the symphysis, spikelets that are
also present in most human mandibles. In man they serve as hitching posts for
the genio-glossal muscles. Von Koenigswald (1956, pp. 111-13) interprets their
presence on the Meganthropus jaw as indicating the power of speech. This inter-
pretation is unfounded, however. See E. L. DuBrul and C. A. Reed: “Skeletal
Evidence of Speech?” AJPA, Vol. 18, No. 2 ( i960), pp. 153-6.
300
The Earliest Hominids
covered Olduvai child. In any case, one cannot expect the jaws
and teeth of related hominids located as far apart as South or
East Africa and Java to be identical, any more than the jaws and
teeth of the human inhabitants of those regions today are identi-
cal.
As to which, if either ( and if not both ) , of these candidates
was our ancestor, no decision can be made on present evidence.
We can state, however, that Meganthropus had not deviated as
far from the human mandibular and dental form as had Zinjan-
thropus.
The Drugstore Australopithecines of China
Because the Djetis fauna is believed to have entered Java
from China, there has been reason to suppose, ever since the dis-
covery of the first Meganthropus jaw, that Australopithecine re-
mains will also turn up in the mother country. In the search of
Chinese pharmacies which turned up the first Gigantopithecus
teeth, von Koenigswald also discovered a few other teeth, which
he tentatively attributed to the Australopithecines, or to a similar
creature, in the following passage. “A few additional teeth that
are not definitely classifiable with either orang or Gigantopithecus
probably indicate the presence of forms related to the Australo-
pithecinae in our fauna. They are of large size, too large for
Sinanthropus, with a very simple cusp pattern, and too small for
Gigantopithecus. These teeth have not yet been studied in de-
tail.” 6
Von Koenigswald sent casts of two canines of this collection to
Broom, who found them similar to those of Sterkfontein and
wrote: “We can, I think, feel fairly confident that a large Aus-
tralopithecine, and probably two, inhabited China in Upper Plio-
cene or Lower and Middle Pleistocene times. And if this should
prove to be the case it may be that they will prove to be even
nearer to man’s ancestor than the South African Australopithe-
6 Von Koenigswald: “Gigantopithecus blacki von Koenigswald, a Giant Fossil
Hominid from the Pleistocene of North China,” Ah AM, Vol. 43 (1952) pp 293-
325-
The Replacement of Australopithecus by Homo 301
cines.” 7 In 1956 Robinson said that these were orangutan canines.8
Who is right I do not know.
The above is the sum total of the data available to me concern-
ing teeth salvaged from Chinese drugstores. Their potential sig-
nificance is enormous.
The Replacement of Australopithecus by Homo
The Meganthropus jaws discovered in Java, the teeth
found in Chinese drugstores, and the hominid fragments unearthed
in Palestine and associated with Oldowan tools suggest that the
Australopithecines spread eastward from Africa across the whole
range of the Old World tropics and inhabited both the Ethiopian
and the Oriental faunal regions. Or they originated in Asia and
spread to Africa. Except for Europe and northern China, Aus-
tralopithecines already occupied, at the end of the Lower Pleisto-
cene, the same territory Homo lived in a little later. In effect, at
the beginning of the Middle Pleistocene Homo replaced Aus-
tralopithecus in the latter’s lebensraum, and his hominid prede-
cessors everywhere vanished from the earth.
The arrival of Australopithecus was almost as sudden as his dis-
appearance. During the last third of the Lower Pleistocene the
bones or tools, or both, of these hominids appear all the way from
Morocco to South Africa, Palestine, Java, and south China. This
is a wide spread for a new genus of primates, which are generally
restricted to single geographical regions at any one time. Just be-
fore their expansion the ancestral Australopithecines must have
acquired some ecological advantage that freed them from local
climatic limitations and afforded them dominance over competing
species.
Tools, a more or less perfect bipedal posture, the ability to col-
lect slow game and to carry it home, the beginnings of a human
type of social structure, a rudimentary kind of hunting, and a
dawning intelligence keener than that of other primates — these
7 Broom and Schepers: “The South African Fossil Ape-Man,” p. 66.
8 Robinson: “The Dentition of the Australopithecinae.”
302
The Earliest Hominids
new acquisitions may have made the Australopithecines more
adaptable to all kinds of tropical environments than were the
forest-bound apes. These are the only traits we know of that
could have provided the needed advantage.
Yet, at the beginning of the Middle Pleistocene, this previously
successful animal faded out rather rapidly and was replaced
everywhere by a different but closely related hominid, man.
Homo must have had an even greater ecological versatility than
his predecessor. What gave him this added advantage? Probably
not stone tools, because the earliest man-made tools are no better
than those attributed to the Australopithecines. Was it fire?
Perhaps, but we have no way of knowing. Only in sheltered
habitation sites, such as caves, can charcoal be expected to survive
the action of wind and water over hundreds of thousands of years,
and when it does we are lucky. Our earliest cave site is Choukou-
tien, in China. The men who lived there 360,000 years ago had
fire. There is no reason to suppose that other men who camped
only in the open did not have it earlier.
In most early habitation sites the broken bones of adult wild
animals bear witness to true hunting. Hunting, speech, fire, and a
type of social organization in which men, under competent leader-
ship and following prearranged plans, could combine forces in
hunts and raids of several days’ duration, must have given Homo
a decided advantage over his less imaginative and less commu-
nicative cousins. At the time he began to replace the Australo-
pithecines he must have possessed such an advantage, or he
would not have won.
Yet so closely similar are the bones and teeth of Australopithe-
cus and Homo that some kind of close genetic continuity be-
tween them must be accepted. But we do not know where or
when the genetic transition from one genus to the other took
place. Did the known Australopithecines, having undergone an
evolutionary sequence of their own, simply become men at the
end of the Lower Pleistocene after they had begun to hunt, to
speak, and to sit around fires; or did some early kind of Australo-
pithecine like Sterkfontein or the Olduvai child evolve into Homo
while Swartkrans and Zinjanthropus pursued their own genetic
blind alleys to extinction? Who knows?
A PRIMATE FAMILY TREE
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304
The Earliest Hominids
The oldest known Australopithecines have been found in Africa,
and the oldest known human remains come from Java. According
to strict chronological sequence, then, a case can be made for the
evolution of the Australopithecines from a local ancestor in East
Africa and their subsequent spread northward and eastward, to
Palestine, south China, and Java. As stated in several papers by
its principal champion, F. Clark Howell, this hypothesis has a cer-
tain logical validity based on a careful scrutiny of the geological
time scales and faunas of different regions.
If we grant the hypothesis and pursue the same logic, then we
may postulate that Homo arose, like Australopithecus, in the
place where his earliest remains are found. But there are two such
places, the Sterkfontein cave and Java. Does this mean that two
races of Homo arose simultaneously from related Australopithe-
cine populations in Africa and Indonesia? Or does it not rather
mean that both Transvaal and Java were marginal areas at the
beginning of the Middle Pleistocene and that, at both, true men
had just arrived from a more centrally located breeding ground
and were in the process of exterminating the Australopithecines?
When Homo and Australopithecus met in such places, as they
probably did, how did they behave toward each other? Did the
men simply hunt down the Australopithecines like impala, or did
they spare some of the females and attempt to mate with them?
Were the two populations mutually fertile, and did some Aus-
tralopithecine genes enter the conquerors’ pools? If so, some of
the regional peculiarities of the earliest men could be easily ex-
plained.
As research workers usually discover at a certain stage of their
inquiries, the more we learn the more complex our problems seem
to become, and possibilities once rejected take on new stature. In
our study of the hominid forerunners of man we have tried to
present every useful scrap of evidence and to examine even the
most remotely possible theories. Whatever the answer is, one fact
is certain. Our present knowledge of the Australopithecines, frag-
mentary and tantalizing as it is, constitutes a most useful back-
ground for the study of the origin and continuity of the races of
man.
a
8
n
AN INTRODUCTION TO
FOSSIL MAN
Of Time, Space, Grades, and Lines
An the last seven chapters I have reviewed some of the
principles of evolution, particularly as they apply to man. I have
described the primates as an order and have traced the descent
of all of them except those that belong clearly to the genus Homo.
Homo can be studied with more insight after this lengthy in-
troduction because now the nature of the scaffolding that holds
up our genealogical structure is discernible. This frame extends in
several dimensions. The first is time, the last half million years or
so, covering the Middle Pleistocene, the Upper Pleistocene, and
the Recent, with one or two possible dips into the tail end of the
Lower Pleistocene. The second is space, which includes the
zoogeographic regions of the world as they exist now and as they
existed during the Pleistocene.
The third is grade, and the fourth is line. Within the dimen-
sions of time and space we have seen several groups of primates
evolve from simple to complex forms and also across one or more
of the biological frontiers that zoologists call adaptive thresholds.
The monkeys of the New and Old Worlds moved from the pro-
simian to the simian grade independently of each other. Each
separately acquired stereoscopic color vision, and in both hemi-
spheres some of them came to brachiate. In the Old World it is
quite possible that at least the latter stages of adaptation for four-
footed life on the ground were reached independently by the
baboons and macaques. The gibbons and the living African apes
An Introduction to Fossil Man
306
also became arboreal separately, as probably did the orang. Thus
three lines of tailless apes independently moved into the pongid
grade. If the hominid grade, with erect posture, was reached by a
single animal only, this border-crossing constitutes a great excep-
tion.
A grade, then, is a stage of physical adaptation to a special way
of life, otherwise known as an ecological niche. A line is a line-
age, a genetic continuum, a succession of animals in process of
phyletic evolution (evolution by succession), from the earliest
distinguishable ancestor to the present form. A line may pass
through several grades and a grade may include populations of
animals belonging to different lines.
Within a line, a population may become extinct in one of three
ways: by dying out completely; by evolving into something
else; or by hybridizing with a genetically different population, and
thus being absorbed.1 All three processes were probably involved
in the evolution of Homo sapiens from H. erectus. The first way is
probably the least important in human evolution because when a
local population dies out, another like it will usually survive else-
where. There is no modern evidence of complete extinction; even
the Tasmanians survive in hybrid form. The second way can be
demonstrated in several lines; and the third, which was probably
the commonest, in many more.
This concept of grades and lines may be the most valuable we
have learned in the last four chapters. In man, as in other pri-
mates, grades and lines are concerned both with ecology and
with anatomy. Among colobine monkeys of Africa and Asia, leaf-
eating is an ecological adaptation for which the stomachs of these
animals have become anatomically suited.
In man the making of tools, the use of fire, and the manufacture
of houses and clothing are all cultural adaptations that provide the
basis of ecological grades, through which, or through some of
which, different human lines have passed. Ecological grades in
human populations involve man’s relationships with the land on
which he lives and with other men. The simplest grade of human
1E. H. Colbert: “Some Paleontological Principles Significant in Human Evolu-
tion,” in W. W. Howells, ed.: Early Man in the Far East (Philadelphia: Am.
Assn. Phys. Anth.; 1949), p. 146.
Of Time, Space, Grades, and Lines 307
culture is collecting wild foods, such as berries, roots, grubs, and
slow game. In essence this is no higher than the ecology of Aus-
tralopithecines, or even of baboons. The next grade is hunting,
which can be further divided technically in terms of weapons and
techniques, such as clubs, simple spears, spears cast with spear
throwers, the bow and arrow, elaborate traps, and the use of dogs.
As previously stated (in Chapter 3), what is important here for
human evolution is not so much the techniques themselves but the
effects they have on human relations and the complexity of social
structure.
After hunting comes agriculture, with or without animal hus-
bandry; specialized pastoral nomadism; village life, with the rise
of arts and crafts; and the births of cities, kingdoms, and empires.
Only the ecological grades preceding agriculture are useful
for present purposes. By the time agriculture began, all the known
subspecies of man had reached their present anatomical forms.
Since then, the racial map of the world has been complicated
and obscured by numerous large-scale migrations and by recent
mixtures. The pertinent cultural grades may be sorted out and
defined in terms of the degree of perfection and diversification of
tools, by whatever conclusions about hunting, food processing,
and skin processing techniques we can draw from studying ani-
mal bones as well as tools in their sites, and by a consideration of
whatever works of art have survived.
In some parts of the world archaeological sequences follow
clear-cut, independent lines through various grades. The most
striking example known of a self-contained cultural line is that of
the American Indians. From a hunting and gathering base with a
technology derived from eastern Asia in the late Pleistocene, they
invented fluted points, polished stone axes, agriculture, pottery,
textiles, bronze metallurgy, urban architecture, writing, and
the concept of zero, apparently independently of the Old World.
Although no other technological line is equally clear-cut and
dramatic, we can trace independent developments, with certain
overlaps, in much earlier periods in several parts of the Old
World. The only line of evidence which we have from the earliest
periods is that of stone tools, including those attributed to the
Australopithecines. The sequence of grades that these tools follow
308
An Introduction to Fossil Man
runs in general from rough core tools to fine core tools, with or
without flakes, to flakes to blades, with or without microliths, to
the use of antler, ivory, and bone, and to the use of polished stone.
In some areas this sequence is incomplete, in others certain stages
have been skipped. In most of them refinements of tool form are
the result of changes in tool-making techniques, from stone-on-
stone to stick-on-stone, to the use of an elastic punch, to pecking,
grinding, and polishing. By studying these technological lines
we can see where contacts between cultures must have taken
place; the diffusion of tool-making techniques from one area to
another indicates communication between different human popu-
lations. As we know from modern examples, this in turn implies
the possibility of gene flow. Where a complete technological dis-
continuity is evident, it generally indicates the arrival of a new
population.
The final application of the concept of grades and lines comes
in the analysis of the bones of the fossil men themselves. Ap-
parently the different parts of the body have evolved to a certain
extent independently of each other and at different rates. The
pelvis and legs, the shoulder girdle and arms, and the skull have
had separate histories. In the following four chapters I will con-
sider the differences in the postcranial skeleton of fossil popula-
tions to the extent that I can, from the grades-and-lines point of
view; but this extent will not be great because the fossil remains
of these bones are few. This subject can therefore be postponed
for the most part to a later volume in which the richer information
on the soft parts of the living will be reviewed.
Here I shall concentrate particularly on the skull, which itself
has a number of components that evolve at different rates: the
fiont teeth (incisors and canines); the cheek teeth (premolars
and molars); the jaws; the brain case; and the mask, meaning the
region of the eyes and nose. From these components, as from the
skull as a whole, it is evident that several lines of fossil men, living
in different regions but within the same time spans, passed
through several cultural grades at different evolutionary rates.
Of the four dimensions time is the least reliable. We do not
know how old all our fossil specimens are, and moreover not all
geologists and physicists agree about the time scale of the second
■
The Dimension of Time 309
half of the Pleistocene, particularly in regard to the correlation of
the glacial and pluvial sequences in different regions. To allot the
remaining chapters of this book on the basis of time would be to
lean the bulk of our weight on the weakest and shakiest timber in
the scaffold.
Space, on the other hand, is the most reliable dimension. We
know exactly where each specimen came from. We also know
something about the distribution of other animals during the
Pleistocene, and in most cases we can relate fossil men to their
faunas. Geography is our strongest timber; it is bolstered, more-
over, by demonstrable sequences of lines of men through various
cultural grades in several regions. In other regions these sequences
are broken, and invasions can be traced both through cultural and
anatomical discontinuities.
The Dimension of Time
In Chapter 7 the chronology of the Lower Pleistocene was
reviewed, primarily as it concerns the places where fossil homi-
nids and tools have been found, that is, Africa, Palestine, and
Java. With the appearance of man in several continents at the
beginning of the Middle Pleistocene the whole world becomes
involved; we must therefore examine the premises on which sev-
eral conflicting evolutionary chronologies are based.
The world is divided into a number of geographical regions,
in which changes in temperature, humidity, soil deposition, and
soil erosion have proceeded at different rates. Regions that lie
near the poles and at high altitudes have proved to be more sensi-
tive to changes in temperature than those that lie near the equa-
tor and at lower altitudes. Equatorial territories, particularly in
Africa, have reflected changes in rainfall more than changes in
temperature. Geologists rely to a certain extent on fauna, which
may change more rapidly in some places than in others. More-
over, we are dealing with an exceedingly brief time span, little
more than a half million years.
The earliest glacial sequence was worked out in the Alps, with
mountain glaciers. Hence the well-known terminology, Giinz,
Mindel, Riss, and Wiirm. However, mountain glaciers are local.
I
310
An Introduction to Fossil Man
and cannot be expected to reflect global climatic changes as
finely as continental icecaps. Of these latter we have three in
Europe — Elster, Saale, and Weichsel — corresponding more or less
to Mindel, Riss, and Wiirm, of the Alpine series. There was no
European icecap to match Giinz. In North America we have four,
Nebraskan, Kansan, Illinoisan, and Wisconsian, which match,
more or less, the four Alpine periods. Like the Alps, the Hima-
layas had four mountain glaciations.
By definition the Middle Pleistocene begins with the advance
of the Elster icecap, Mindel I, and the Second Himalayan Glacia-
tion. It does not include Giinz or the First Himalayan, which have
been relegated to the very end of the Lower Pleistocene, along
with the Cromerian Interglacial. Before the Villafranchian was
cut loose from the Pliocene and added to the bottom of the
Pleistocene, the situation was much simpler; the Pleistocene was
simply the Ice Age. These new complications make the problem
of dating the phases of the Middle and Late Pleistocene even
more difficult.
Penck and Bruckner, who worked out the Alpine sequence, set
the beginning of Giinz 600,000 years ago, and that of Mindel
about 500,000 years ago. They allotted 60,000 years for the Giinz-
Mindel Interglacial, 240,000 for the Mindel-Riss, and 60,000 for
the Riss-Wiirm. According to their scheme, Wiirm lasted 60,000
years and ended sometimes between 16,000 and 24,000 years ago.
We know now that it ended about 10,000 years ago. As 10,000 plus
60,000 equals 70,000, their corrected date for the beginning of
Wiirm is about 70,000 years ago. This has recently been more or
less confirmed by a Carbon- 14 date of 64,000 ± 1,100 years ago
for a mild climatic oscillation shortly after the beginning of
Wiirm I.2
This is the earliest Carbon-14 date yet determined anywhere.
It sets the outer boundary of our ability to date sites by this well-
known method.3
2 H. Godwin: “Carbon-Dating Conference at Groningen” (September 14-19,
1959), Nature, Vol. 184, No. 4696 (1959), pp. 1365-6. The C-14 date number
is GRO-1379.
3 On January 10, 1961, the National Bureau of Standards announced that the
half life of Carbon-14 is now 5.760 years instead of 5.568, as formerly believed.
The old dates were calculated on the former basis. They must now be multiplied
The Dimension of Time 311
Penck and Bruckner made their calculations by standard geo-
logical procedure, including measurements of rates of erosion,
and estimates of the rates at which soils were deposited on land
surfaces. Aside from the Wiirm, they did not try to date the dura-
tion of the glaciations, only that of the interglacials and of the
Ice Age as a whole. As this procedure has been followed by
most geologists, it is difficult to find one who will commit himself
on the lengths of the periods of glaciation and thus construct a
complete glacial chronology.
Zeuner, however, did suggest a complete glacial chronology,
using all data available in 1951, including calculations based on
the amount of solar radiation that reached various latitudes of
both the Northern and the Southern Hemisphere of the earth’s
surface at different times. These figures, which a modern elec-
tronic computer could deliver in a few hours, were the result of
twenty years’ efforts by the Yugoslav M. Milankovitch and his
associates. The computations are based on the correlations of three
astronomical cycles: (1) changes in the angle between the equa-
torial plane of the earth and the plane of its orbit (40,000 years);
(2) variations in the season at which either hemisphere passes
closest to the sun in the course of orbit (92,000 years); and
(3) the periodicity of a slight conical movement in the earth’s
axis (26,000 years). Unfortunately these correlations are so
complicated that they may be interpreted in more than one way.
At any rate, with this and other aids, Zeuner filled in the 600,-
000 year period of Penck and Bruckner as follows. Mindel began
about 500,000 years ago and reached two peaks, 476,000 and
435>000 years ago. The Mindel-Riss Interglacial lasted 190,000
years; Riss reached two peaks, 230,000 and 187,000 years ago.
The Riss-Wiirm Interglacial lasted 60,000 years; and Wiirm
by 1.0345. In most cases, the difference falls within the range of probable error
of the sample, and the maximum difference is about 2,000 years. The reader may
make these corrections if he wishes.
Following procedure initiated by E. S. Deevey of Yale in 1961, I have desig-
nated each Carbon-14 date by its laboratory number. For example, GRO-1379 is the
date given in footnote 2. GRO means Groningen, Netherlands. Other symbols used
are NZ for New Zealand, W for Washington, L for the Lamont Laboratory of Co-
lumbia University, C for Chicago, P for Philadelphia (University of Pennsylvania),
RM for the Rritish Museum (Natural History), and I (UW) for Isotopes Inc New
York.
I
312
An Introduction to Fossil Man
reached three peaks, 115,000, 72,000, and 25,000 or 22,000 years
ago, depending on the latitude. Since the end of Wiirm III, 22,000
years have elapsed.4
Zeuner admits the possibility of an error of 20 per cent for
Wiirm III, and the C-14 process indicates that his dates for
Wiirm are on the whole too early. However, he believes that the
further back one goes in time the smaller the error, and he allows
a deviation of 5 per cent for the earlier glaciations.
In 1946, Harold C. Urey discovered that, as water evaporates,
the three isotopes of oxygen, oxygen 16, 17, and 18, go off at
slightly different rates, and the lightest of the three, oxygen 16,
goes off the most rapidly. He then studied the proportions of
oxygen isotopes in the carbonate deposits, formed mostly of
foraminifera, at the bottom of the ocean. After many refinements,
he and his associates developed a method for dating long cores of
carbonates drilled from the ocean’s floor.
Intensive work on this material was carried out in the early
1950’s, particularly by Cesare Emiliani, who finally developed a
sequence that shortened Penck’s and Bruckner’s estimate of the
Ice Age by about a half. According to Emiliani, Giinz (Nebras-
kan) extended, in round numbers, from 300,000 to 265,000 b.p.
(before present); Mindel (Kansan) from about 200,000 to 175,-
000; Riss ( Illinoisan ) from 125,000 to about 100,000; and Wiirm
( Wisconsian) from 70,000 to 10,000. The sea-water isotope system
gives Wiirm only two subperiods, known as stadials, pinpointed
at 71,000 to 57,000 and 28,000 to 8,ooo.5
More recently, Rhodes W. Fair bridge has worked out another
timetable based on the fluctuations in sea level along the various
shores of the earth. This, too, is very complex and involves a num-
ber of factors and variations in different regions. He finds that,
although the interglacials were warmer, the mean sea levels were
100 meters higher than today before the glaciers began to ac-
cumulate; that at the height of the Giinz-Nebraskan, despite the
accumulation of ice at the poles, the shores were still 30 meters
above present level; that at the peak of the Mindel-Kansan they
4F. Zeuner: Dating the Past, Third Edition (London: Methuen & Co.; 1952),
pp. 144-5-
5 C. Emiliani: “Ancient Temperatures,” SA, Vol. 198, No. 2 (1958), pp. 54-6.
The Dimension of Time 313
were virtually at present sea level; during the Riss-Illinoisan they
went down to 87 meters below present sea level; and the two
peaks of Wiirm-Wisconsian found them to be between 85 and
100 meters below the present level.
Fairbridge, by correlating his recent sea-level dates with the
Carbon- 14 record, has confirmed the accuracy of his method in
detail for the post-Wiirm oscillations. Extending it backward, he
finds a general agreement with Emiliani’s data, but stretches the
dates out a little; he would put the peaks of Gimz-Nebraskan at
320,000; those of Mindel-Kansan at 230,000; of Riss at 112,000;
and Wiirm at 62,000 and 25,000 b.p. Like Zeuner, he correlates
his findings with the solar cycles of Milankovitch, but with a dif-
ferent result. If he is correct, the Middle Pleistocene began 250,-
000, instead of 500,000, years ago. As the beginning of the
Pleistocene is still placed at about one million years ago, the span
of the Lower Pleistocene would then be about 660,000 years,
two thirds of the Pleistocene rather than one half of it.
While these ingenious calculations were being made, teams of
physicists in Germany and California experimented with a more
long-ranged method, the measurement of Argon-40.6 Argon-40
and Calcium-40 are both formed by the decay of Potassium-40.
When a crystalline mineral containing Potassium-40 is heated to
500° F. or more, the Argon-40 which had previously been formed
inside it, and which had been trapped, escapes. After the mineral
cools, new Argon-40 atoms collect inside it.
Such a mineral is anorthoclase, one of the feldspars. Anortho-
clase is ejected onto the earth’s surface by volcanic eruptions, and
when it emerges it is too hot to contain Argon-40. As time goes on,
this gas accumulates in it at a fixed rate. The measurement of the
proportion of the gas in the mineral tells, within a probable error
of five to seven per cent, exactly how long ago the volcano
erupted. Argon-40 is also formed in tektites, which are small glassy
meteoric nodules found in many countries. As they are heated
while passing through the earth’s atmosphere, Argon-40 begins to
6R. W. Fairbridge: “The Changing Level of the Sea,” SA, Vol. 202, No. 5
( i960), pp. 70-9.
G. H. Curtis: “A Clock for the Ages: Potassium Argon,” NG, Vol. 120, No. 4
(1961), pp. 590-2.
314 An Introduction to Fossil Man
form in them after they have landed and cooled, just as it does in
lava.
Not only have these physicists dated deposits in Olduvai Gorge,
Tanganyika, and the Trinil Beds of Java (as stated in Chapter 7),
but they have also given us several critical dates for glacial events
in Europe and North America.7 The early glacial till of California,
equivalent to the local Donau glaciation in Central Europe,
which was pre-Giinz, was dated at 850,000 b.c. This makes sense
in terms of the date of one million years ago for the beginning of
the Villafranchian. Giinz is moved back to 500,000 years ago, and
Mindel to 400,000. The latter part of the Second, or Great, Inter-
glacial is set at 230,000 years, a date based on samples taken from
the Late Acheulian site at Torre in Pietra, Italy. Hand axes from
this site are similar to those found with the Swanscombe skull in
England. This is as close to the present as Argon-40 had taken us
early in 1962, and it leaves a gap of 160,000 years to the oldest
Carbon-14 date. On the whole it ties in well with the older system
of Penck and Bruckner and of Milankovitch, and fails to support
those of Emiliani and Fairbridge. The last two seem to be more
accurate for the latter part of the Pleistocene than for its earlier
part.
Fairbridge’s work sheds light on another subject — the avail-
ability of land bridges at different periods. For example, the pas-
sage across Bering Strait, where the sea-level fluctuations have
been simple, was possible only during the peak of the Riss-Illinoi-
san, and all of the Wiirm. Fairbridge’s view also reduces the pos-
sibility that the Djetis fauna, about which much has already
been said, could have reached Java very much before the begin-
ning of the Middle Pleistocene. This is heartening to paleontolo-
gists and archaeologists as it indicates that fewer early sites lie
lost forever under the sea than we had feared.
These new dating methods and correlations unfortunately do
not greatly concern Africa, where so much evidence of early
7J- F. Evemden, G. H. Curtis, and R. Kistler: “Potassium-Argon Dating of
Pleistocene Volcanics,” Quaternaria, Vol. 4 (1957), pp. 13-18.
W. Getner and J. Dahrinder: “The Potassium- Argon Dates of Some Tektites,”
ZfNF, Vol. 14a, No. 7 ( 1959), pp. 686-7.
K. P. Oakley: “Dating the Stages of Hominid Evolution,” The Leech, Vol. 28,
Nos. 3, 4, 5 (1958), pp. 112-5.
The Dimension of Time 315
hominid evolution has been found. There anthropologists work
with a series of pluvials ( wet periods ) and interpluvials ( dry pe-
riods) originally aligned as follows: Kageran Pluvial = Giinz; Ka-
masian Pluvial =Mindel; Kanjeran Pluvial = Riss; and Gamblian
Pluvial = Wiirm.
Unfortunately the Kanjeran Pluvial has been clearly identified
only in East Africa. In other parts of the continent south of the
Sahara only three pluvials are known. This discrepancy has led
some geologists to believe that the Kanjeran is only a subdivision
of the Kamasian Pluvial, which reached one or more peaks just as
in Europe each of the four glaciations did. At any rate, the end
of the Kanjeran, whatever its status, is clearly marked, for it was
then that the faulting that cracked open the Rift Valley and left
it in its present form is believed to have taken place.
Determining whether sub-Saharan Africa underwent three or
four pluvial periods during the Pleistocene is only part of the
problem, for the periods of wetness and drought varied in differ-
ent parts of the subcontinent, just as today some parts are wet
and others dry. In the past, zones of moist climate, with their ap-
propriate floras and faunas, shrank and expanded gradually, so
that at any one time a particular spot on the map could have been
wet while another a few hundred miles away was dry, and a little
later, geologically speaking, both could have been either wet or
dry. The climate of South Africa has been particularly deviant
because it has been affected by air masses from Antarctica which
follow a pattern of their own independent of that of the Northern
Hemisphere. This complicates the problem of establishing a date
for the Australopithecines and for the human-looking jaw frag-
ment, Telanthropus 2, from Sterkfontein Cave.
In North Africa and in the Near East the Pleistocene sequence
is based partly on changes in fauna and partly on a complex and
imperfectly understood series of local rises and falls of the sea
levels along both the Atlantic and the Mediterranean coasts.
In Java,8 which like Africa has been found to contain both Aus-
tralopithecine and very early human specimens, geological re-
search on the Pleistocene has been concentrated in the Solo River
Valley in the eastern part of the island. There deep cuts through
8 H. R. Van Heekeren: “The Stone Age of Indonesia,” VKIV, Vol. 21 (1957).
316
An Introduction to Fossil Man
successive layers show sequences as long as those in Olduvai
Gorge, and like Olduvai’s they are composed in part of volcanic
materials potentially suitable for argon-potassium dating.
The Pleistocene is represented by three superimposed beds,
which are, starting at the bottom, the Putjangan, Kabuli, and
Notopuro. These contain, in the same order, the Djetis, Trinil,
and Notopuro faunas. The first two are more or less continuous
geologically and in fauna, but the Notopuro beds are distinctly
marked. Just before they were formed the land rose, creating
new drainage lines. At the same time Sundaland emerged from
the sea during what corresponded to the Third, or Riss, Glacia-
tion. At the end of the Notopuro period the waters again rose, and
the succeeding fauna is modern.
The age of the Djetis fauna is still in dispute, but this fauna
probably existed in the Late Lower Pleistocene, corresponding to
the Cromerian Interglacial in Europe. The Trinil fauna is Middle
Pleistocene, and it lasted until the ocean level fell again in the
Third, or Riss, Glaciation. The Notopuro fauna is Upper Pleisto-
cene. As we shall see in Chapter 10, three distinct but successive
forms of man inhabited Java during these three divisions of the
Pleistocene, but it is impossible for us to pinpoint their dates with
greater accuracy.
In China 9 also only major divisions of the Pleistocene are read-
ily discernible because that country was unglaciated east of the
Tibetan Highlands. China is divided into two geological regions,
north and south, by the ridge of the Tsinling Mountains. This
range runs east and west in Shensi Province, on about the 34 ° lati-
tude, just south and west of the first great bend of the Huang Ho.
In north China the Pliocene-Pleistocene threshold is marked by
a series of warpings and faultings in the earth’s crust, followed by
the deposition of the Nihowan-Taiku, or Horse Beds, laid down
in lake bottoms and river channels. They contain a local Villa-
franchian fauna known as the Sanmenian series, which includes
horses, elephants, cattle, sheep, deer, and camels. The climate
was cool.
The Lower Pleistocene of south China is harder to identify. In
9 Cheng Te-Kun: Archaeology in China, Vol. l of Prehistoric China, Heffer,
Cambridge, 1959.
The Dimension of Time 317
Kwangsi, whence most of the material comes, the beds are later-
ites (red soils produced by rock decay) washed from the valley
slopes and deposited in terraces 12 meters above the present
high-water levels. The fauna is the same as that of north China.
In north China the Middle Pleistocene deposits are called
Terra Rossa, because they consist of red conglomerates and red
clays. In south China a second set of terraces and river fans com-
posed of water-borne laterites represents this period. At this time
and in both regions many caves and fissures were opened and
then filled up with air-borne earth. This earth then solidified, im-
prisoning large numbers of animal bones, including those of
Sinanthropus and a somewhat later human skeleton, that of the
Ting-tsun man, both of which will be described in Chapter 10.
Throughout the Middle Pleistocene north China had a Pale-
arctic fauna although the climate varied intermittently from cool
and semi-arid to almost tropical. In south China, where the entire
period was tropical, the fauna was Indo-Malayan, as described
in Chapter 7. The fossil remains of this fauna come mostly from
the yellow deposits of the Kwangsi caves.
As the deposition of Terra Rossa soils continued without a
break, the Upper Pleistocene arrived in north China unostenta-
tiously, and the only way in which geologists can tell the Early
Upper Pleistocene layers from the Late Middle Pleistocene ones
is by the fauna. No new species appeared, but some of the old
ones had become extinct by the Upper Pleistocene.
Then in the middle of the Upper Pleistocene, the land rose
again in another continental uplift, which was followed by the so-
called Chingshui Erosion. After that, yellow earth was deposited
in Kansu, Shensi, and Shansi by northwest winds, which also
brought a cool and semi-arid climate. This was the Age of Yellow
Earth, equivalent to the Wurm glaciation in Europe. Its fauna was
the same as before, but further impoverished by continued extinc-
tions. In south China this period is unknown. At the end of the
Pleistocene came the Age of Black Earth, whose soils contained
a transitional fauna between the fauna of the Yellow Earth and
the modern animals of China.
In the two Americas and in Australia, continents remote from
the centers of human evolution, human beings arrived so late that
An Introduction to Fossil Man
318
we have little need of reviewing geological details in this book.
All finds are well within the time span of the Carbon- 14 clock.
More exact intercontinental correlations than those outlined
here may be expected when the Chinese begin dating their sites
by Carbon- 14 and when the Argon-40 method shall have been
more widely used. More generally, as the Space Age reaps the
fruits of the Atomic Ages, these chronological problems may be
solved within our lifetime. But they have not been solved yet, and
several interesting fossils have been set aside by scientists because
their exact age is unknown. Time is still not our most reliable
yardstick.
The Dimension of Space: Glacial Geography 1
For present purposes the dimension of space consists of the
geography of the land masses of the earth during the last half
million years, particularly the areas of land covered by ice, the
continental shelves exposed by the drying of oceans, and the
intercontinental land bridges that afforded animals and men tem-
porary passage between zoogeographic regions.
During the glacial maxima, ice covered most of northern and
eastern Europe, some of the Tibetan plateau, and parts of the
diagonal mountain spine of Central Asia. It nearly sealed off
Western Europe from Eastern Europe and it blocked passage be-
tween China and the West, except for those hardy animals able to
negotiate chilly passes in summer. Until at least the Third Inter-
glacial it is fairly certain that man was not one of them.
If Fairbridge is correct, both the Sunda and the Sahul shelves
could have been dry land during the Riss glaciation and also dur-
ing the Wiirm. Multiple and separate invasions of Indonesia from
southeast Asia would then have been possible, and even Australia
could theoretically have been invaded by man more than once.
Bering Strait, which is only 200 feet deep, could have permitted
the crossing of ancestral Indians from Siberia to Alaska during the
Riss-Illinoisan glaciation, if any such ancestors were on hand at
1 The most useful general source is J. K. Charlesworth: The Quaternary Era
(London: Edward Arnold, Ltd.; 1937).
THE WCiRM GLACIATION IN EUROPE
AND CONTEMPORARY SEA LEVELS
320
An Introduction to Fossil Man
that time. During the Wiirm-Wisconsian the passage was wide
open, and before the end of that period America was inhabited by
its basic aboriginal population.
During most of the time which concerns us Great Britain was
part of the European continent. The Strait of Gibraltar, now 1,000
feet deep, was an open-water barrier throughout this period. Those
who have crossed between Spain and Morocco in small craft know
that the tides, currents, and winds can make this passage danger-
ous. The sides of the cut through which the water flows are very
steep. Even when the sea level was 300 feet lower, something
sturdier than rafts or simple canoes would have been needed to
mount an invasion mustering more than a handful of people.
Although no certain evidence exists to prove it, possibly a land
bridge connected Tunisia and Italy during part of the Lower
Pleistocene, allowing sabertooths and a few other African ani-
mals to pass into Europe.2 Also, momentarily during the Upper
Pleistocene, narrow channels between islands in this part of the
Mediterranean may have permitted human passage, but not an
exchange of fauna.
During glacial maxima the Caspian Sea rose high above its pres-
ent level, up to 300 feet during Riss and 250 feet during Wiirm.
At the times of flooding the Caspian waters, fed by the Volga and
by the glaciers that it drained, flowed across the strip of lowland
north of the Caucasus to spill into the Black Sea. But the Black
Sea had its high water when the Caspian was low, and vice versa,
because only during interglacials, when the oceans were high and
the Mediterranean swollen with salt water, did the Mediterranean
floods break through the Bosporus to fill the Black Sea basin.
Between these periods the Black Sea was a brackish lake. The
Aral Sea was enlarged in rhythm with the Caspian, and south and
east of the Urals stretched a vast swamp, below the edge of the
ice. Europe was as difficult to approach from the northeast as
from Africa. Its only gateway to the outside was the Bosporus
and the Levant.
During pluvial periods similar floodings took place in Africa,
in what is now the Sudan and southern Sahara. Lake Tchad, now
a shrinking body of shallow water, was once a broad lake. To its
2 Charlesworth: op. cit., pp. 1226-7.
321
The Dimension of Space: Glacial Geography
east extended, at least intermittently during the Pleistocene, an
extensive area of swamps and sometimes possibly of lakes. This
barrier extended from the Sabaluka Gorge, 50 miles north of
Khartum, some 450 miles southward to about 10° North Latitude.3
These water barriers, and the existing great lakes of East Africa,
which were greatly enlarged during pluvials, must have restricted
animal and human traffic in Africa, moving both north and south
and east and west, to a few narrow highways, and made Black
Africa nearly as inaccessible as western Europe. But after the
Pleistocene the lakes and swamps shrank and the East African
highlands were invaded at least twice by people from the north.
For the Lower Pleistocene the faunas of different regions serve
as fair indicators of the passage of time because evolution was
then working overtime. For the Middle and Upper Pleistocene
the chief value of faunas lies in their record of extinctions. Time
is measured by the number of species that had disappeared at
each period, with a few exceptions. The spotted hyaena, Crocuta
crocuta, first appeared early in Mindel, more or less simultane-
ously in Europe, Africa, and Asia, and his presence serves to cor-
relate these continents chronologically.
There is also a geographical aspect to faunal distribution dur-
ing the Pleistocene. In Europe the Middle Pleistocene is marked
by the arrival of a new set of animals, the cold fauna, all of which
— the mammoth, woolly rhinoceros, reindeer, and others — had
become cold-adapted by means mentioned in Chapter 2. They
replaced the warm or Villafranchian fauna of the Lower Pleisto-
cene, one member of which, the hippopotamus, continued to live
in his chilly rivers until the beginning of the Elster glaciation.
The cold-adapted animals of the European Middle Pleistocene
were Palearctic and were more closely related to Oriental than
to Ethiopian species. North African mammals were Ethiopian al-
most until the end of the Pleistocene, when Palearctic species ap-
peared, including the bear, stag, and European wild boar (Sus
scrofa ), and also the European elk (moose to Americans), whose
portrait has been found among the rock paintings of the Sahara.
In southeast Asia three successive faunas migrated, as we have
3 G. Andrew: “Geology of the Sudan,” Chapter 6 of J. D. Tothill: Agriculture
in the Sudan (Oxford: Oxford University Press; 1948), pp. 84-128.
322
An Introduction to Fossil Man
already discussed, from India and China into Indonesia. These
movements are important to the subject of this book, because
where edible animals go, man the hunter follows.
The Temporal and Spatial Distribution of Fossil Man Sites
At the time of writing at least 337 sites which can be dated
with some degree of accuracy have yielded skeletal remains of
TABLE 1 1
FOSSIL-MAN SITES IN TIME AND SPACE
Time
Region
L&M
IG3
W-l
W-2,3
P-W
Total
Per Cent
of Total
Western Europe
3
6
29
78
37
153
49
Eastern Europe and
U.S.S.R.
0
2
6
20
5
33
10.6
North Africa
1
4
1
2
14
22
7.1
Africa south of Sahara
1
2
3
1
15
22
7.1
Near East
0
1
10
6
4
21
6.7
India and Ceylon
0
0
0
0
2
2
.6
East Asia
5
5
0
2
2
14
4.5
Southeast Asia and
Indonesia
4
1
1
1
17
24
7.7
Australia and New
Guinea
0
0
0
1
3
4
1.3
America
0
0
0
4
13
17
5.4
Total
14
21
50
115
112
312
100.0%
Per Cent of Total
4.5
6.7
16.0
36.8
35.9
99.9%
Time Symbols: L&M = Lower and Middle Pleistocene
IG3 = Third Interglacial
W-l = Wtirm 1
W-2,3 = Wtirm 2 and 3
P-W = Post-Wiirm
The Temporal and Spatial Distribution 323
fossil men that reputable scientists have recorded.4 These bones
represent a minimum of a little over one thousand individuals,
ranging in completeness from a tooth to a skeleton. These 337
sites are distributed in space, and with various degrees of prob-
ability in time, as follows.
In the time scale on Table 11, the first column is labeled
Lower and Middle Pleistocene” for the benefit of the possibly
human Telanthropus mandibles from Sterkfontein and of the
Pithecanthropus specimens from the Djetis faunal beds of Java.
There is still some doubt whether all these remains belong to the
Late Lower or Early Middle Pleistocene. The rest of the sites in
this column are unquestionably of Middle Pleistocene date.
In any case, this portion of the Pleistocene ( 85 per cent, more
or less), from our standpoint the most important since it was the
formative period for H. erectus and H. sapiens , is represented by
only fourteen known sites, or 4 per cent of the whole. In only one
of them, Choukoutien, were there more than a few fragments of
one, or at the most three, individuals. The earlier half of the
Upper Pleistocene, consisting of the Riss-Wiirm, or Third, Inter-
glacial in Europe and the Kanjeran-Gamblian Interpluvial in
much of Africa, is represented by only twenty-one sites, or 7 per
cent of the whole, although some other, imperfectly dated pieces
might be included. The second half of the Upper Pleistocene, that
is, Wiirm, Wisconsian, or Gamblian, can also be divided, this time
into periods of no more than 35,000 years each, covering the
consecutive regimes of Neanderthal and Upper Paleolithic men
in Europe and the arrival of human beings in Australia and the
Americas. The first of these periods is represented by fifty sites, or
16 per cent of the whole; the second by 115 sites, or 37 per cent.
Into the few thousand years between the last retreat of the
Scandinavian icecap and the diffusion of agriculture must be
crowded 112 sites, or 36 per cent of the total.
It is easy to read into these figures an increase in the human
population in Pleistocene and early post-Wiirmian times similar
to the increase currently taking place, but such an interpretation
must be made with caution. Long after the beginning of human
4 V. Vallois and H. L. Movius, Jr.: Catalogue des Hommes Fossiles (Al-
giers, 1952); and reports of other sites published since this compilation.
An Introduction to Fossil Man
324
existence, the dead were still being abandoned, to be bitten,
crunched, and dismembered like the bodies of any other crea-
tures. Some living tribes continue this custom to this day.
Deliberate burial did not begin until the Late Pleistocene, and
has never been universally practiced. Short of mummification, the
best way to preserve (and unwittingly ensure the discovery of)
a skeleton is to bury the body in a cave, and this was not done in
many parts of the world. In any case, it was not done anywhere
before the Late Pleistocene. Burial customs, or their absence,
probably affect the numbers of skeletal specimens discovered in
different periods much more so than population size. However, the
world’s population undoubtedly grew slowly as new regions were
settled and new techniques of food acquisition invented.
We must also consider the geography of archaeological search.
Paleolithic archaeology was born in France. The French have
many archaeologists, much limestone, and many caves. More than
34 per cent of the world’s known sites containing human remains
are in France or in present or former French colonies or de-
pendencies. Even outside these territories the French have been
active. More than 22 per cent of the sites are in British Common-
wealth territories, for the British have been almost equally en-
thusiastic. Three per cent are in former or present Dutch terri-
tories, which encompass only a small part of the land area of the
world; and the Dutch have found five of fourteen Lower and
Middle Pleistocene sites. Were the count made by the nationality
of the discoverers, the French, British, Dutch, Germans, and
Americans would be far in the lead, for between them they have
found well over 90 per cent of all fossil-man sites and specimens.
Time , Space, and Paleolithic Tools
Before we discuss ways and means of studying the thousand-
odd human fossils of the Pleistocene listed in the previous sec-
tion, it may be useful for us to study the distribution of Paleolithic
tools, for two reasons.
Tools are more abundant than human bones. An adult human
body has only 180 or so bones, many of which contain edible sub-
Time, Space, and Paleolithic Tools 325
stances, brain or marrow. Paleolithic stone implements are made
of very hard materials, including quartz, quartzite, chert, chal-
cedony, and obsidian. During his lifetime a hunter makes thou-
sands of implements that are inedible and almost as incorruptible
as gold. As indicators of the presence of man, stone tools are more
useful than human bones.
Moreover, stone tools constitute the principal source of informa-
tion about the cultural life of Pleistocene peoples. Tool-making
techniques are handed down from generation to generation, and
sequences of such techniques indicate cultural lines. When two
groups of people whose territories have common borders are
found to have made similar tools, we may infer that one group
taught the techniques to the other, with an added likelihood of
gene flow between them. Conversely, when two lines of tool-
making follow similar evolutionary paths, although they are
widely separated in space, the possibility of independent inven-
tion and parallel cultural change must be considered.
The study of Paleolithic tools also has drawbacks. The earliest
tools must have been so crude that they are indistinguishable
from naturally fractured stones. Endless arguments have taken
place concerning the identification of eoliths, or dawn stones, as
these dubious specimens are called. In addition, we do not know
whether the oldest tools were made by men or by Australopithe-
cines. In one site, Olduvai Gorge, Australopithecine bones were
found in association with stone tools on a floor on which the
Australopithecines bad lived. Both these problems were discussed
in Chapter 7. During the Lower Pleistocene both Australopithecus
and Homo may have made tools, but from the beginning of the
Middle Pleistocene onward, all the tools we have were probably
made by Homo.
Aside from choppers and chopping tools, which have already
been described, these tools fall into four principal classes: bifacial
hand axes, flakes, blades, and microliths. Detailed descriptions of
these are readily available and need not be repeated here.5 In
general, bifacial hand axes are usually large, almond-shaped im-
plements flaked on both sides and both bilaterally and bifacially
5 Oakley: Man the Toolmaker (London: Brit. Mus. Nat. Hist.).
C. S. Coon: The Seven Caves (New York: Alfred A. Knopf; 1957), pp. 29-41.
326
An Introduction to Fossil Man
C
HAND AX
Fig. 43 Basic Tools of Early Men. A. Chopper, from Melville Island, Australia
( Tiwi ) ; B. Chopping tool, from Melville Island, Australia (Tiwi); C. Hand ax,
from England ( Seven Caves ) ; D. Simple flake tool, from Le Moustier, France
(after Oakley, 1956); E. Levallois flake tool from Syria; F. Blade tool from Syria.
The earliest tools made by man were choppers, which have a cutting edge flaked
on one side only (A) and chopping tools, flaked on both sides (B). They were
made from Africa to China in the Late Lower Pleistocene. In east and southeast
Asia they persisted through the Pleistocene, and in parts of Australia were made
in the twentieth century. Hand axes (C) were confined to Africa, Europe, and
southwest Asia as far east as India. Simple flake tools were made wherever tools
were used. The one shown here (D) was made by a Western Neanderthal. Flake
tools made by striking prepared cores were also used by Neanderthal men, par-
ticularly in southwest Asia ( E ) . Blade tools, characteristic of the Upper Paleolithic
of Wiirm II, had a limited distribution from England to Afghanistan.
Time, Space, and Paleolithic Tools
symmetrical. They may be shaped in either of two ways: the hand
axes proper are pointed at the business end, and the cleavers have
transverse, bladelike edges in place of points. The hand axe was
probably an all-purpose cutting tool whereas the cleaver may have
been somewhat specialized, as for skinning animals, felling small
trees, or both.
Flake tools are classified by the technique used to strike the
flakes off the parent core and by the treatment given the flakes
after they have been detached. The crude way to make flakes con-
sists first of trimming the chalky crust off the surface of the core
by a series of glancing blows, leaving the core polyhedral rather
than rounded; then searching the surface for a place where two
planes form a relatively sharp angle and striking the core with a
hammerstone just above that ridge, so that a more or less tri-
angular flake will spring loose at a single blow. Unless the flake
is unusually well shaped it will need trimming at the butt and
along the edges before it can be used as a knife, scraper, spear
point, or whatever.
Another relatively crude method consists of setting the core on
an anvil stone and striking it on top with a hammerstone, so that
flakes fly off both ends and all sides. Experts can distinguish these
flakes, which are called bipolar, and also their cores.
But if the tool-maker knows how, he can shape the core in ad-
vance by knocking off a small chip here and another there,
thereby visualizing the flake that will come off. In particular, he
fashions a striking platform with an angle as close as possible to
45° so that he will get a nearly flat flake. When he strikes the
critical blow, if all goes well, his flake will fall off, ready for use.
This third technique produces what is known as a Levallois
flake.
Once a flake has been detached, it can be used as it is, and
perhaps sharpened after it has suffered a few nicks; or it can be
trimmed at once into a special shape. Sharpening and trimming
are called retouching. Its butt may be trimmed to thin it for
hafting, or its edges and point retouched to make it into a side-
scraper or end-scraper. A sharp blow at the tip will turn it into
a narrow chisel or graver. These extra preparations suit it for spe-
cial work with wood, skins, flesh, bone, antler, or ivory. In some
r
An Introduction to Fossil Man
328
of the most advanced flake cultures, as in the Near East and
Europe, specialized flake tools are numerous.
Blades differ from flakes in that they are essentially parallel-
edged instead of triangular, and usually thinner and straighter.
They are elongated strips of flint or obsidian made by a special
process, as follows. The tool-maker skillfully shapes his core so
that it is virtually tubular, with a flat top serving as a striking
platform. Near the edge of this flat surface he sets the point of a
punch made of an elastic material such as fresh bone, antler, or
even wood, and holds the punch at right angles to the plane of
the striking platform and directly in line with the main axis of the
core. Striking carefully placed blow after blow, he slivers off blade
after blade. These blades are the blanks from which he can re-
touch a wide variety of specialized tools.
The fourth step in tool-making is to prepare small cores in the
manner just described and then strike off miniature blades that
can then be retouched into microliths — fine tools such as tiny
gravers, or small, individual blades set in rows inside slots carved
in wooden or bone handles. Such rows of microblades are known
as composite tools, which take the forms of knives, sickles, or
barbed spear and arrow points.
On the basis of the five types of tools described here and in
Chapter 7, the most anciently inhabited parts of the Old World-
excluding Oceania and Australia — may be divided into two ma-
jor provinces: a Western, situated in Europe, Africa, and West
Asia, and an Eastern, in East Asia and Indonesia. During the
entire Middle and most of the Upper Pleistocene, the borders of
these two provinces met only in India. Thus the ancient homes
of the Australoids and Mongoloids lay in the Eastern Province,
and those of the Caucasoids and Africans — Capoids and Con-
goids — in the Western.
In the northern region of the Eastern Province the tool types
were derived directly from those of the Lower Pleistocene and
evolved independently throughout the Middle and Upper Pleisto-
cene. Gradually the choppers dropped out of the sequence. At
first both simple and bipolar flakes were made, but the bipolar
ones also dropped out and the sequence as a whole was essentially
Time, Space, and Paleolithic Tools 329
a gradual refinement of the simple flake techniques plus an in-
creasing use of antler and bone. During the Upper Pleistocene
Western-style flakes and blades appeared at different times on the
western and northern fringes of the Eastern Province, but neither
of them altered the evolutionary course of tool-making in the
province as a whole.
In Europe, western Asia, and most of Africa, hand axes ap-
peared at the beginning of the Middle Pleistocene, first crudely
and later finely made. The cruder ones are called Abbevillian or
Chellian; the finer ones, Acheulian. The Abbevillian style lasted
until the beginning of the Mindel-Riss Interglacial, and the
Acheulian disappeared as an industry early in the Riss-Wiirm In-
terglacial, although special kinds of hand axes were made, as rare
tools, almost until the First Wiirm Interstadial,6 some 40,000 years
ago. In Africa the rate of change was slower, and in the Cape
Province an Acheulian industry was replaced by a flake industry
as late as 30,000 years ago. The sequence in India does not mir-
ror those of Europe and Africa exactly, and it is not certain when
hand axes ceased to be made there.
From the beginning of the Middle Pleistocene flake tools co-
existed with hand axes, either as elements in hand-axe cultures
or as cultures of their own.7 Only in the late Upper Pleistocene
did flake tools entirely replace them. Flake tools were most elab-
orate and specialized in the Near East and Europe, where three
partly successive and partly overlapping flake cultures filled the
time span from the beginning of the Middle Pleistocene to the
end of the Early Wiirm, some 40,000 years ago, and lasted
sporadically even later than that.
These were the Clactonian, a simple flake industry that in-
volved crude retouching; the Levalloisian, which included pre-
pared cores and Levallois flakes; and the Mousterian, a simple
flake industry that involved fine retouching. In some places the
Levalloisian and Mousterian techniques were combined into a
hybrid culture known as Levalloisio-Mousterian. In North Africa
6 An interstadial is a mild, or cool, period between two peaks of a glacial pe-
riod. It is shorter and less warm than an interglacial.
7 The word culture is used here in the special archaeological sense.
330
An Introduction to Fossil Man
a local flake culture, the Aterian, followed the Acheulian and de-
veloped into a highly specialized, technically refined way of pro-
ducing knives and projectile points finely retouched on both
sides. Some were tanged, barbed arrowheads strikingly similar to
later American Indian specimens.
In the deep forests of Central and West Africa the tradition of
the hand axe continued, in a special industry called the San-
goan, well into the Late Pleistocene and perhaps almost until its
end. In East and South Africa flakes of the general Levalloisio-
Mousterian type replaced Acheulian hand axes progressively, the
transition working its way generally from north to south, until,
as stated above, it took place in Cape Province more than 100,000
years later than it did in the Near East and Europe.
Blade-making was invented in the region of Palestine, Leba-
non, and Syria, or nearby, during the Early Wiirm, and this new
technique was apparently brought to Europe by way of the
Bosporous gap in the warm Gottweig, or Wiirm I — II, Interstadial.
By 30,000 b.c. Upper Paleolithic hunters who made both blade
and flake tools reached the Atlantic, and over the next 20,000
years a bewildering (to me) sequence of Upper Paleolithic tool
industries followed. Microliths were part of the European tool
kit from the start. To the East blade-making extended as far as
northern Afghanistan, where C-14 dates as old as those in west-
ern Europe have been determined.
From their European and central Asiatic centers blade-making
techniques were carried northeastward into Siberia and over the
mountains and narrow channels of icy water to Hokkaido, where
a number of presumably successive blade cultures has been found.
Just before and after the end of the Pleistocene, blade cultures
were brought to North Africa in two waves, the Mouillian and
Capsian, and there these new techniques replaced the Aterian
flake industry. Capsian tool-making was also carried to East Af-
rica in early Recent time, and it eventually spread to South Africa
in a mostly microlithic culture known as Wilton.
In southeast Asia and Indonesia the earliest tools were chop-
pers and chopping tools, with crude flakes. These have been found
in Burma, Malaya, Siam, Indochina, Luzon, Borneo, Java, and
Sumatra. Only in Malaya have they been definitely dated — at
Time, Space, and Paleolithic Tools 331
the local equivalent of the Cromerian Interglacial or the begin-
ning of Mindel I. This was the time of the Djetis fauna in Java.
As in China, the flake tools in these industries gradually evolved
into finer forms, followed in postglacial times by local microlithic
industries.
In Australia, which was uninhabited until almost the end of
the Pleistocene, all types of tools were either imported from In-
donesia or invented locally. During the last century, and in some
places during the twentieth, tool industries had a continent-wide
distribution. Choppers as crude as any known in the early Middle
Pleistocene were used until very recently (and perhaps still are),
on Mornington Island in the Gulf of Carpenteria, as they were
two generations ago on Melville Island. Good Levallois-like flakes
were made in central Australia and blades and microliths in Vic-
toria and neighboring parts of New South Wales. In several re-
gions aborigines had learned to grind chopping tools to a smooth,
sharp edge and to haft them as axes with sticks and gum. Ar-
chaeologically these cultures go back to a C-14 date of 6740 b.c..
The New World industries stem from two sources, the Late
Pleistocene flake culture of China, and the blade cultures of
northeast Siberia, which had originally come from the West. In
America these imports evolved into a number of industries some
of which lasted until the arrival of Europeans. Very crude chop-
ping tools have been collected in both North and South America,
and they are as crude as the tools of Sinanthropus. We do not
know how old they are, although we may soon. In South America
they are found in beds underlying pottery, and were probably
used by some of the Fuegian Indians until quite recently. The
presence of these archaic tools cannot be taken as proof that
Homo erectus preceded Homo sapiens in America.
The foregoing archaeological summary indicates that the world
of the Middle and Upper Pleistocene was divided into two halves
and five regions; an eastern province with a northern and a south-
ern region, and a western province with a Eurasian, a North Af-
rican, and a sub-Saharan region. These correspond, as we shall see
shortly, to the distribution of the five human subspecies near the
end of the Pleistocene, before the movements that relocated the
332
An Introduction to Fossil Man
Australoids and Capoids when the two dominant northern sub-
species, the Mongoloid and the Caucasoid, expanded their terri-
tories southward.
The Chronology and Distribution of the Use of Fire
Much has already been said in this book about the impor-
tance of fire to the physical and cultural evolution of man, and it
need not be repeated here. But if it can be shown that some geo-
graphical races got fire before others did, the implication will be
that those who had it first were also the first to receive its evolu-
tionary benefits, and that those who obtained it last must have
been correspondingly retarded.
Unfortunately, the absence of fire can be indicated only by
negative evidence. We cannot expect to find charred wood and
bone in disturbed sites such as gravel beds, and if we find such
evidence, as found it was at Swanscombe on the Thames, we are
extremely lucky. In Java we would not expect to find it for we
have no undisturbed sites there. Only in Africa is there evidence
that fire arrived late, as late as 40,000 years ago. In the earlier
habitation sites such as Olorgesailie in Kenya, where layer after
successive layer of hand axes, cleavers, and meat bones have
been excavated with the most meticulous care, not a trace of
charcoal or charred bone has been found.
Both Louis Leakey and Desmond Clark, who are among the
most painstaking and observant excavators in the world, have
stated their conviction that in East Africa the entire hand-axe
period was fireless almost to the end. If future excavations con-
firm this erudite opinion, we shall have one explanation of the ex-
traordinarily slow pace that human evolution followed, in the
Middle and Late Pleistocene, in Africa south of the equator, and
perhaps also south of the Sahara.
Grades and Species of Fossil Men
During the Pleistocene, hominids made tools in five traditions.
As far as we know at present, tool-making began in Africa, in the
333
Grades and Species of Fossil Men
second half of the Lower Pleistocene, with split pebbles, chop-
pers, and chopping tools. This simple technology spread as far as
southeast Asia and Indonesia. At this point the hominid world
knew but a single way of making tools.
Before the Lower Pleistocene was over, and perhaps even be-
fore the original tool-making tradition had reached southeast Asia,
the tool-makers of Africa and Palestine had added coarsely
chipped, ball-like implements to their repertoire, and these new
items were apparently not diffused to the East. Thus the split be-
tween East and West, of which Kipling sang a half million years
later, opened before the end of the Lower Pleistocene. In the
West, where hand axes were invented a little later, a further divi-
sion into three parts did not occur until the end of the Middle
Pleistocene. By that time the Chinese and southeast Asian in-
dustries had also become recognizably different from each other.
If the first tools in each region were made by Australopithecines,
and we have no evidence to the contrary, we may infer that the
first split mentioned above took place between separate popula-
tions of Australopithecines belonging either to two or more spe-
cies, or to two or more subspecies. But at the dawn of the Middle
Pleistocene, when Australopithecus and Homo coexisted in Java
and probably also in Africa, the same kinds of tools were ap-
parently being made by men who, as the millennia rolled on, in-
vented new ways of working with flint and other tool materials
and new and more efficient tool forms.
From this evidence a second inference is logical, that men
evolved from Australopithecines in either or both provinces. No
more than two such evolutionary acts are implied by existing
information. The only alternative is the theory that Homo and
Australopithecus lived side by side during the Lower Pleistocene,
that only Homo made tools, and that in both Africa and east Asia
he preyed off his hominid cousins until he had destroyed them.
This second hypothesis is not impossible but it is unsupported.
As long as we use the word Australopithecine in a broad sense,
including forms undiscovered as well as those known, we may
postulate without serious reservation that Homo was descended
from a polytypic species of Australopithecus inhabiting a wide
stretch of Old World tropics. By the same logic we may, as others
334
An Introduction to Fossil Man
have done, speak of an Australopithecine grade of human evolu-
tion.
A grade is not a formal taxonomic unit, like family, subfamily,
genus, species, or even waagenon (see Chapter 1, p. 17). It is an
evolutionary stage or condition, as broad or as narrow as circum-
stances require. For example, as we saw in Chapters 4 and 6, all
the New World primates and most of the Old World genera
passed independently from a prosimian to a simian grade, and
three lines of Old World simians passed separately to the pongid
grade. Although also called a subfamily, the Dryopithecines con-
stituted a grade through which more than one line passed. Some
of these lines became independently specialized into hominids
and pongids during the latter part of the Miocene or the Pliocene.
We can therefore tentatively set up, in our family tree, a suc-
cession of three grades: Dryopithecine, Australopithecine, and
Hominine. The only professionals who may be expected to object
to this scheme, which is not original, are the anatomists and
paleontologists who believe that man is descended from Oreo-
pithecus, and those others who believe that Homo’s ancestors
parted from those of Australopithecus by evolution through
branching before the latter had evolved into their known forms.
The second objection is largely one of nomenclature or semantics,
and depends to a certain extent on the forthcoming study of the
Fort Ternan maxilla and teeth; on whether the Olduvai child
was a true Australopithecine, a Hominine, or an unnamed ances-
tor of both; on a detailed study of the Tchad skull; and on the
reservations we may have concerning the new finds that may
await us in the ground.
Once past the Australopithecine stage, we come to the full Mid-
dle Pleistocene. Homo is already divided into a number of geo-
graphical populations. How many grades, from that point on, shall
we recognize in fossil and living men?
Some authors prefer three, variously labelled. A compromise
nomenclature is Protoanthropic, Paleanthropic, and Neanthropic*
8 This classification is derived from S. Sergi’s categories Protoantropi, Palean-
tropi, and Fanerantropi , and from J. Piveteau’s Archanthropiens, Paleanthropiens ,
and NJanthropiens. The term Archanthropi would make them to men what arch-
angels are to angels; and Faner&ntropi, from phaneros, Greek for visible, is un-
familiar.
GRADES AND LINES OF FOSSIL HOMINIDS
HS = Homo sapiens HE = Homo erectus A = Australopithecine
Fig. 44
An Introduction to Fossil Man
336
The first grade includes such obviously ancient and primitive
fossils as Pithecanthropus, Sinanthropus, and the new Chellian-3
skull from Olduvai Gorge. The second covers a mixed group — the
European and Near Eastern Neanderthals, Solo in Java, and
Broken Hill in Africa — which have little in common except long
low skulls, big brow ridges, and an Upper Pleistocene date. The
third includes modern man, of all races, and all fossil men that
could be more or less duplicated among the living.
This threefold system is unrealistic: its grades are partly based
on the time scale, and they should be entirely dependent on size,
which is not considered, and form. Ignoring time, we shall see that
the small-brained Solo and Broken Hill skulls belong to the first of
these grades, and that other members of the second grade can be
matched, in most respects, among living peoples. The second
grade, then, dissolves, and the third requires subdivision.
There remain two mutually exclusive grades, the Protoanthropic
and Neanthropic. The Paleanthropic does not represent a true
unit; its members can be sorted and redistributed in the other two
categories. These two differ sufficiently in several pertinent re-
spects to warrant being given the status of separate, successive
species, Homo erectus and Homo sapiens. For those who like to
split categories more finely, each species can be divided into
grades of lesser magnitude.
Between some of the fossil populations of our own species,
Homo sapiens , anthropologists have observed what seem to be im-
portant differences in evolutionary status. But when viewed from
the perspective of life as a whole, these variations are slight. Dif-
ferences of equal magnitude still exist. Bij definition, every minor
grade discernible in fossil specimens of Homo sapiens may be
found in living men.
The great variability of twentieth-century human beings, in
evolutionary grades as well as in racial lines, makes our lives more
complicated than those of our ancestors who lived in a simpler
world, when space was still a barrier to communication and travel,
S. Sergi: “I Tipi Umani Piu Antiche,” Chapter 3, pp. 69-133 of R. Biassutti:
Razze e Popoli della Terra (Turin: Union Tipografico-Editrice; 1959), Vol. 1.
J. Piveteau: Traite de Paleontologie, Vol. 7 of Primates, Paleontologie Humaine
(Paris: Masson et Cie; 1957).
The Sapiens-Erectus Threshold 337
and local populations kept to themselves. As we move rapidly
around the world, these differences pose a challenge that we may
or may not be sapient enough to meet.
The Sapiens-Erectus Threshold: the Evidence of Brain Size
In order to decide whether a given human fossil specimen
belongs in the category of Homo erectus or that of Homo sapiens,
we need as much evidence as we can get. Usually, however, very
little evidence is available, and for practical purposes it is most
often limited to the skull. Of the fossil skulls known to us only
nineteen are so different from those of living peoples that they
obviously belong in the erectus category. They are, counting
adult skulls only: three Pithecanthropus specimens from Java;
six Solo skulls from the same island; six Sinanthropus skulls; one
newly found skull from Tze-Yang, China; and, from Africa, the
newly found Chellian-3 skull from Olduvai Gorge, the Saldanha
Bay skull, and Broken Hill ( Rhodesian ) man.
These skulls have in common small brains, low cranial vaults,
heavy brow ridges, and sloping foreheads. The palates and teeth
that are preserved are big; and the available lower jaws that
match these skulls lack chins.
Certain other fossil skulls, like the Upper Paleolithic crania
from Europe and the Upper Cave family from Choukoutien, are
entirely modern. In one way or another most of the other skulls
and groups of skulls are modem only in the sense that the primi-
tive features which they possess may also be seen in the crania of
surviving primitive peoples. This intermediate group is sapiens in
the Linnean sense, that is, that all living men are sapiens.
With these facts in mind, just where do we draw the line? Ob-
viously, the differences between the two species, one of which
evolved by succession out of the other, are concerned with intelli-
gence, self-control, and the abilities to provide food efficiently
and to get along well in groups. The seat of intelligence is the
central nervous system. The regulation of self-control is the com-
bined task of the brain and the endocrine system, and indeed the
brain and endocrines act together in many ways, and influence
each other by a complex feedback system.
338
An Introduction to Fossil Man
Fig. 45 Anatomy of the Skull. A. Side view; B. Front view, of a South African
Bantu skull; C. Interior basal view of a laboratory skull; D. Section through the
sella turcica region of same. Abbreviations: CF. Cerebella fossa; E. Ethmoid bone;
ES. Ethmoidal sinus; F. Frontal bone; FM. Foramen magnum; L. Lachrymal bone;
Mn. Mandible; Mx. Maxillary bone; N. Nasal bone; O. Occipital bone; P. Parietal
bone; PI. Palatal bone; S. Sphenoid bone; ST. Sella turcica; T. Temporal bone;
Z. Zygomatic bones ( malar ) . ( Drawings A and B after de Quatrefages and Harny,
1882; C and D after Hamilton, 1956. )
The size of the human brain is related to a capacity for per-
formance in thinking, planning, communicating, and behaving
in groups, as leader, follower, or both. But brain size is a sum of
the masses of that organ’s component parts, including the me-
dulla, hypothalamus, cerebral hemispheres, and cerebellum. The
The Sapiens-Erectus Threshold 339
hemispheres are divided into lobes, and their surfaces are covered
with a wrinkled skin of gray matter, the cortex, which contains
neurones. In living individuals and populations, differences are
found in the relative sizes of the lobes and in the surface areas of
the cortex; the size of the surface area varies with the complexity
and depths of the folds on the inner and outer surfaces of the
hemispheres. The larger a brain is, the greater the cortical surface
area, both proportionately and absolutely. The cerebellum is a
miniature replica of the greater part of each hemisphere, and it is
covered by 75 per cent as much cortical surface, but its cortex is
thinner. As the hemispheres grew larger in the course of human
evolution, the cerebellum expanded proportionately.
As in the case of the Australopithecines, all that remains of the
brains of fossil men are imprints left on the inside of the skull.
These show gross size and form fairly accurately, and a few other
details as well. The divisions between the lobes are marked in
varying degree, and the seat of the cerebellum in the basal part of
the occipital bone may be detected as a pair of cups. The blood
supply to the covering of the brain may also be gauged to a certain
extent by the imprints of the middle meningeal arteries on the
parietal bones.
The inside of the skull base also contains the sella turcica
(Turkish saddle), also known as the hypophyseal fossa. This is a
depression, or cradle, in the midline of the sphenoid bone and ex-
tending to either side. In it the pituitary gland, or hypophysis, is
seated. The length and depth of this depression is taken to be a
rough indication of the size of the pituitary — the master gland
which, among many other functions, controls growth, including
that of the bony crests that brace the skull. No other endocrine
gland leaves a direct mark on the skeleton.
In mammals brain size, like the face length of horses ( see Chap-
ter x, page 25), is allometric. That is, in any closely related group
of animals, such as a family or subfamily, all species of which are
more or less equal in intelligence, brain weight will equal body
weight times a standard fraction and carried to a given power.
This formula expresses the principle that the large species will
have absolutely larger but relatively smaller brains than the small
species.
340
An Introduction to Fossil Man
However, groups that differ in intelligence (more properly
known as level of cerebral evolution ) have different formulae.
And as we go up the scale, from marsupials, for example, to pro-
simians to monkeys, apes, and finally men, these formulae grow
less and less accurate. As Jerison has discovered,9 in the most
highly evolved animals brain weight acts not as a single unit but
as a sum of two units, only one of which varies allometrically as
expected. The other is less influenced by body size, if at all.
The first unit, which in man constitutes only 10 to 12 per cent of
the total, corresponds to virtually the total brain weight of a prim-
itive mammal, such as an opossum. The second unit, on which
human intelligence primarily depends, cannot vary in a simple,
allometnc way: whether a man is large or small, he needs a certain
number of neurones and their connecting fibers to enable him to
behave like an intelligent human being.
If a fossil man of modern body weight had a cranial capacity
two thirds that of the modern range ( assuming brain weight and
cranial capacity to be roughly equivalent), the second unit of his
brain weight would have been no more than 82 per cent of the
whole. Therefore the differences in intelligence between Homo
erectus and Homo sapiens were presumably greater than a gross
comparison of brain sizes would indicate.
Also, in comparing individuals, we need not be greatly con-
cerned with differences in brain size due to body size. In living
men the allometric (or, one might say, body-weight sensitive)
proportion of brain weight varies from about 150 grams in a 100-
pound person to about 225 grams in a 150-pounder. This maxi-
mum difference of 75 grams is important principally in comparing
male and female skulls.
The nineteen fossil skulls which are morphologically clearly dif-
ferentiated from those of living persons have cranial capacities
ranging from 775 cc. to 1,280 cc. Most modern skulls range from
1,200 cc. to 1,800 cc. The only two real series of erectus skulls that
we have, those of Sinanthropus and Solo, which have six skulls
each, range from 1,035 to T255 cc. (the figures are identical), and
both have means of 1,095 cc. The only one of the other seven skulls
®H- J- Jerison: “Brain to Body Ratios and the Evolution of Intellijr
Science, Vol. 121, No. 3144 ( 1955), pp. 447-9.
ence,”
The Evidence of Cranial Form 341
which exceeds this range is that of Broken Hill, Rhodesia, with
1,280 cc. Because Broken Hill man was a large male, the 25 cc.
excess could easily have been due to allometry.
The smallest skull that is morphologically excluded from the
erectus category is also the oldest one that can be called sapiens.
It is the Steinheim specimen, a small female cranium which has
not yet been thoroughly studied and which needs reconstruction
because it is crushed. Its capacity is variously given as anywhere
from 1,170 to 1,290 cc. If the Steinheim woman weighed as little
as 90 pounds, which is possible, she can be allowed an allometric
deduction of 50 cc. in brain size in comparison to the others, and
this places her near the top of the erectus brain-size range, or even
over it. In any case, she is within the modem female range.
The approximate threshold between the brain-size ranges of
Homo erectus and Homo sapiens can then be set at about 1,250
to 1,300 cc., with the expectation that some of the fossil sapiens
skulls will, like many modern ones, be smaller. The designation of
a fossil skull as erectus or sapiens depends on the total configura-
tion, and not on brain size alone.
The Evidence of Cranial Form
In general. Homo erectus crania are long and broad at the
base and converge toward the top, both lengthwise and sidewise.
Homo sapiens skulls bulge more in front and back and at the
sides, because a larger brain is set on a proportionately smaller
base. Conventional measurements designed for the living are less
useful in describing these differences than a few special measure-
ments taken on individual bones of the skulls of both species,
erectus and sapiens, particularly in the sagittal line of the frontal,
parietal, and occipital bones. Two principal measurements are
taken on each bone: its external sagittal arc, i.e., its length meas-
ured along the center line of the skull; and its chord, i.e., the dis-
tance, in a straight line, between its two ends.
In the nineteen skulls considered to be erectus, the frontal is the
longest of the three bones and the parietal the shortest. In modern
men the parietal is characteristically the longest and the occipital
11
342 An Introduction to Fossil Man
the shortest. In some of the Neanderthals, which are excluded
from the erectus category on the basis of brain size and some
other features, the frontal bone may be longer than the parietal
because the lengths of the brow ridges, which are heavy, are ordi-
narily included in frontal length.
BREGMA
Fie;. 46 Sagittal Arcs and Chords in Homo erectus and Homo sapiens. One
of the most reliable methods of distinguishing the two species of Homo is by com-
paring the three sets of arcs and chords of the skull in sagittal section. Homo
sapiens is more curved in the frontal and parietal segments than Homo
erectus; the opposite is true of the occipital segment.
For each of the three bones a sagittal curvature index may be
computed by dividing the chord length times 100 by the length of
the arc. The frontal bone has two curves, one extending from the
343
The Evidence of Cranial Form
root of the nose (nasion) over the crest of the brow ridges to the
base of the forehead proper (glabellare) and a second from the
base of the forehead to bregma — the place where the frontal and
parietal bones meet at the crown of the skull. If the frontal bone
is treated as a whole, these two curves tend to cancel each other
out, so that no difference is seen between H. erectus and modern
men. If, however, separate indices are calculated for the two parts
of the frontal bone, the glabellar (brow ridge) part has an index
of from about 75 to 85 in the H. erectus group, whereas in modern
man this index runs from about 85 to nearly 100. In the other part
of the frontal bone, the forehead section, the figures are nearly
reversed, because modern men have relatively curved foreheads.
The index of curvature of the parietal bone is a better criterion,
because the sagittal profile of this bone forms a simple arc. In the
skulls called erectus in this study, the parietal chord is 93 to 97 per
cent of the length of the arc; in modern skulls it runs from about
89 to 92 per cent. Little or no overlap in this ratio may be found
between the successive species. In the occipital arc-chord index,
the chord is 73 to 76 per cent of the arc length in the erectus skulls,
as compared to a range of 79 to 86 per cent in Homo sapiens.
Whereas the figures for erectus skulls indicate a true range, those
for sapiens skulls indicate instead a range of means (statistical
averages) — no figures for a range of individual sapiens skulls are
obtainable. For this reason the two sets of figures are not quite
comparable and the amount of overlapping between the two spe-
cies in these indices is unknown.
These interspecific differences reflect principally the increase in
brain height which occurred during human evolution, and the
reduction of bony crests which took place as the frontal lobes grew
over the eye sockets, rendering special protection above the eyes
unnecessary, while the occipital crests were needed less and less
as the head achieved a more perfect balance on the cervical verte-
brae.
These arc-chord indices may be used along with absolute brain
size as an additional set of criteria in sorting fossil skulls into the
two species. Many other criteria are possible, including face
breadth, forehead breadth, the dimensions of the bony eye sock-
344
An Introduction to Fossil Man
ets, and those of the cranial base, but because faces and bases are
usually missing or defective in the oldest skulls, these constants
are of less value than the three vault indices described above.1
The Evidence of Tooth Size
Contrary to what a number of textbooks say, we cannot
make a blanket statement that human teeth have grown progres-
sively smaller in the course of evolution, because this is true of
some populations but not of others. Tooth reduction has moved at
different rates in different lines. In certain lines, like that of the
Bushmen, a rapid reduction can be traced, and in others, like that
of the Australian aborigines, teeth are almost as big as ever. The
size range in modern human teeth is very great but modern teeth
as large as most of those of Homo erectus are exceptional.
We have 147 Sinanthropus teeth representing about 32 indi-
viduals, 20 Pithecanthropus teeth from at least three individuals,
and 16 from the single Broken Hill cranium. Lower teeth only are
also available from four mandibles found in North Africa: three
at Ternefine in Algeria and one at Sidi Abd er-Rahman in Mo-
rocco. All four are old enough and morphologically suited to
qualify as Homo erectus, if only tentatively.
With three exceptions, all the teeth listed above fall within the
size ranges of the teeth of modern men, either in crown length or
crown breadth, or in both dimensions. The three exceptions,
which have both longer and broader crowns than any correspond-
ing modern teeth, are the following: an upper canine in the palate
of Pithecanthropus 4, from the Djetis beds of Java; a lower second
premolar from the Pithecanthropus B mandibular fragment,
found in the same beds; and another lower second premolar im-
bedded in mandible No. 2 from Ternefine. And that is all. No cor-
responding teeth attributed to Homo sapiens in this book are
known to be as large as these three.
1 R. Martin and K. Sailer: Lehrbuch der Anthropologie, Third edition (Stutt-
gart: G. Fischer; 1957-1961), Vol. I, 1957, and Section 8, 1959.
F. Weidenreich: “The Skull of Sinanthropus pekinensis,” PSNSD, No. 10
(i943), PP- 128-31.
G. Schwalbe: “Uber die Beziehungen zwischen Innenform und Aussenform
des Schadels,” DAKM, Vol. 73 ( 1902), pp. 359-408 (for the frontal indices).
A Brain-Size to Tooth-Size Index
345
A Brain-Size to Tooth-Size Index
Tooth size is not the best criterion of human evolution. But
teeth have, in a very general way, grown smaller while brains have
been growing larger, and an index that mirrors these movements
in opposite directions is more sensitive to change than either com-
ponent. Two such indices can be made: a brain-palate index,
already mentioned in connection with the Australopithecines,2
and a brain— molar-size index devised for this occasion.3
TABLE 12
THE BRAIN-PALATE INDEX AND
THE BRAIN-MOLAR INDEX
Brain-Palate Index Brain-Molar Index
Age-
Rank
Subject
Index
(3) Zinjanthropus 1.1
(2) Swartkrans 1.2
(1) Sterkfontein 1.4
(4) Pithecanthropus 4 1.4
(5) Sinanthropus 1.7
(9) Broken Hill 1.7
(11) Tasmanian (mod.) 1.8
(7) Gibraltar I 1.9
(8) La Chapelle 1.9
(6) Steinheim 2.0
(10) Combe Capelle 2.2
(11) Modern English 2.3
Australo-
pithecus
II. erectus
H. sapiens
Age-
Rank
Subject
Index
(3) Zinjanthropus 22
(2) Swartkrans 22
(1) Sterkfontein 24
(4) Pithecanthropus 4 30
(5) Sinanthropus 34
(10) Broken Hill 37
(6)
(8)
(9)
(7)
(12)
(11)
Steinheim
Skhul V
Combe Capelle
La Ferrassie
Cro-Magnon
Grimaldi
39
40
40
43
43
43
hn Table 12 the first two grades of each list are as inclusive as
I could make them. However, the third grade, that of Homo
sapiens, represents a selection, because of the enormous amount of
calculations necessary to cover the entire field. In general, the
columns follow the same order, from Australopithecus to Homo
erectus to Homo sapiens. The Homo sapiens sections contain only
This is a modified version of Sir Arthur Keith s index, explained on page 292 n.
The brain-molar index is the cube root of cranial capacity divided by 100
times the square root of the sum of the crown areas ( length x breadth ) of the six
upper (when possible) molar teeth. Or:
I = vAc/^S 1. X b. (M1, M2, M3) X 100
346
An Introduction to Fossil Man
European and Near Eastern material except for the modern Tas-
manians in the brain-palate index. They are listed to show that
evolution proceeds at different rates in different lines, as the pres-
ence of Broken Hill man in the erectus category also indicates.
Had we enough patience, ingenuity, and time, it would be easy
to invent dozens of other indices and ratios to finish staking out
the erectus-sapiens frontier, but we have not. What has been pre-
sented will have to suffice. The moment has come to move on to a
matter of almost equal theoretical importance, the further evolu-
tion of man once he had crossed the border into Homo sapiens.
Evolutionary Changes within Homo Sapiens:
the Rise of the Chin
I f b y Homo sapiens one thinks principally of living Europeans,
and in particular of the articulated skeletons of urban paupers
dangling from hooks in European and American lecture rooms,
then indeed many differences may be found between some fos-
sils that in this book have been labelled Homo sapiens and the
mounted specimens just mentioned. But if we compare fossil men
such as the Neanderthals with the peripheral, primitive popula-
tions of the world, the gap between living and fossil sapiens skele-
tons narrows, until it is closed. Brow ridges reach their peak on
Melville Island, and mastoids their minimum in South Africa.
Chief among the hallmarks of the sapiens state in the works of
many writers is the presence of a chin, despite the fact that chins
have turned up in Sugrivapithecus (one of the Indian god-apes),
Kromdraai, and gibbons. Many who sneer at phrenology believe
that a prominent chin is a badge of courage, firmness, and deci-
sion. Consciously or unconsciously, those who make the chin a
sine qua non of being Homo sapiens have fallen into a lexical trap.
In English, French, and several other languages, the word mental
means both “of or pertaining to the chin” (mention), and “of or
pertaining to the mind” (mens) [Webster], We have no evi-
dence that chins and I. Q.’s have anything in common, or that a
Cro-Magnon could outwit a Neanderthal by virtue of his mental
protuberance. The absence of a chin does not exclude any indi-
vidual, living or dead, from the species Homo sapiens.
Evolutionary Changes within Homo Sapiens 347
In fossil sequences modern-style chins appear toward the end
of each phyletic line, starting at about 30,000 b.c. in Europe and
somewhat later elsewhere. If we postulate that the chin arose
through a single mutation which had to be spread by migration
and mixture, or by migration and replacement, we have the Up-
per Paleolithic Europeans moving over the earth at nearly jet-age
speed, and violate the evidence of geography and cultural history.
Chins obviously appeared in each population, if and when
needed, in response to forces of a mechanical nature. Reduction in
tooth size was an influential, but far from the only, factor. We
know this because many modern peoples have teeth as large as
those of their chinless ancestors. For example, the teeth of Heidel-
berg man, well preserved in the chinless Mauer mandible, are no
larger than those of Upper Paleolithic Europeans.
According to E. L. DuBrul and his associates,4 who have spe-
cialized in this subject, the chin seems to have been formed in
response to a combination of separate but related changes in three
organs or sets of organs : the four pairs of muscles that principally
operate the lower jaw — temporals, masseters, lateral pterygoids,
and medial pterygoids; the teeth; and the tongue, with its set of
governing muscles. As the amount of chewing needed for survival
diminished, the size of these four pairs of muscles could be re-
duced. A man could live out his life safely, from the nutritional
point of view, with smaller teeth than his ancestors had. With
smaller muscles and smaller teeth there was no further need for
the mandible to remain large and massive, and, along with the
palate, it eventually became smaller, and in particular, thinner
and more delicate.
This general reduction in mandible and palate size produced
crises for the medial pterygoid muscles and for the tongue. Each
median pterygoid muscle is attached at one end to an area on the
inside of the gonial angle of the mandible, and at the other end to
a point on the palate just behind the third upper molar. As the
rear attachments are farther apart than the forward attachments
4 E. L. DuBrul and H. Sicher: The Adaptive Chin (Springfield, 111.: Charles C
Thomas; 1954).
DuBrul: Evolution of the Speech Apparatus (Springfield, 111.: Charles C
Thomas; 1958).
An Introduction to Fossil Man
348
of these muscles, each time the pair of them is contracted as a unit,
in order to draw the mandible back in chewing, strain is put on
the center of the mandible, which must be strong; otherwise the
two halves of the bone would break apart. In the process of evolu-
tion the front attachments of the medial pterygoid muscles mi-
grated to the rear as the whole masticatory apparatus was re-
Fig. 47 The Lateral Pterygoid Muscles and the Chin. Of the five principal
sets of paired muscles that move the lower jaw, the lateral (external) pterygoids
have a special function which is related to the evolution of the chin. These
muscles move the mandible forward in rotary chewing. They extend from the
condyles of the mandible to the palatal bone, just behind the upper third molar
teeth When the jaws are at rest, these muscles are extended (A). But when the
mandible is moved forward, they contract ( B ) . Because the forward anchors of the
muscles on the palatal bone lie inward from the condyles as well as forward,
when the muscles are contracted (B) they pull the condyle inward as well as for-
ward If the jawbone were not very strong at its symphysis, it would snap like a
wishbone. The shorter the mandible the stronger its symphysis, everything else
being equal. When, in the evolutionary process, the jaw became shorter and the
basin in which the tongue and other neighboring organs rest was lowered, a
strong brace was still needed. That brace moved outward, with the formation' of
new bone (the mental trigonum), and became the chin. (Drawings after DuBrul
and Sicher, 1954.)
duced, whereas the rear attachments remained in the same
positions as before. This change widened the angle between the
two muscles and increased the strain on the mid-line of the bone
per unit of force exerted by the muscles themselves. As the func-
tion of these muscles is to move the jaw backward, there is no
reason for it to have become reduced to the same extent as did the
muscles that clamp the jaws together. Therefore the reduction in
muscular effort exerted by the medial pterygoids was largely com-
pensated for by the increase in strain on the chin area per unit
of force.
Evolutionary Changes within Homo Sapiens 349
In chinless primates this strain is taken by a brace on the inside
of the mandible, in the form either of a general thickening or of a
bar known as the simian shelf. In Homo erectus there is no simian
shelf, but the bone is thick. Whereas the outer surface of the man-
dible is smoothly convex at this point, the inside is concavo-con-
vex, and sloping strongly inward.
In a very large jaw, the presence of this brace permits space
within the oral cavity for a tongue and its guiding muscles of the
same size as those of living people. If the size of the mandible
were to be reduced and its shape remain constant, then the
tongue would be badly crowded, especially as the palate also be-
came shorter and shallower. Therefore, in the course of the jaw
reduction that took place in human evolution, the shape of the
mandible had to change. Room had to be made for the tongue.
The lower borders of the mandible moved outward, and the cen-
tral brace migrated from the inside to the outside of the center
line, producing the human chin.5
The tongue itself could not be reduced in size along with the
teeth and jaws because we need it for talking, and we talk more
than we swallow. This does not mean that the appearance of a
chin made speech possible, only that it kept speech from becom-
ing difficult as the jaws grew smaller. The absence of a chin in
Homo erectus does not tell us whether or not he could speak, or,
if he could, when he began to do so. Only through a careful study
of the nervous system can these questions be answered, and in
Homo erectus such a study is difficult if not impossible.
A reduction in the amount of chewing also affected some of the
cranial bones. As the temporal muscles shrank, the zygomatic
arches under which they move grew less flaring and the face be-
came narrower. Thick skull bones and heavy braces, including the
brow ridges, were no longer useful for survival as men invented
more efficient ways of killing than clubbing people over the head.
With (or possibly without) a change in endocrine balance, skulls
6 It is easy to test this hypothesis. For example, I have measured the cubic
capacities of shellacked casts of the Heidelberg and Cro-Magnon mandibles. Both
are large. The former is chinless; the latter has a prominent chin. To the bottom
and rear of each specimen I attached a cut-out floor of waxed cardboard, and
then I filled each with water up to the level of the tooth line. Each contained 5.5
fluid ounces, or about 163 cc.
35°
An Introduction to Fossil Man
Fig. 48 How Brow Ridges Protect the Eyes against Blows. In Homo erectus
(above), the eye sockets lie in front of the brain case, and brow ridges are needed
to protect the eyes from blows. In Homo sapiens (below), the eye sockets lie under
the brain case, which affords the eyes the same protection. This figure also il-
lustrates the difference in brain shape, in the lateral view, between Homo
erectus ( based on Pithecanthropus 4 ) and Homo sapiens ( a brachycranial Aus-
trian skull ) . ( Drawings after Moss and Young, 1960. Their drawing is based on
Weidenreich: The Skull of Sinanthropus, Fig. 270 B & C.)
could become thinner and brow ridges smaller by the usual
biological process that prunes organs and structures which are no
longer needed. Like tooth reduction, these changes take time, and
some living populations are closer to Homo erectus in these re-
spects than are others.
Lines and Subspecies of Fossil Men: the Evidence of Teeth
I f w e could find a few specimens of Homo erectus and Homo
sapiens frozen in the ice of the Mindel glaciation, we might learn
Lines and Subspecies of Fossil Men 351
how ancient are the modern variations in skin color, beard and
body haii development, and hair form. But as such a windfall is
less than likely, we must content ourselves with the racially vari-
able parts of the human body which are available. These parts,
bones and teeth, indicate that from the very beginning the men of
the Lower Pleistocene, whether called erectus or sapiens, dif-
fered from each other regionally in features and anatomical de-
tails that can still be recognized in living men. Among them are
details of tooth anatomy.
In addition to variability, durability, and abundance, teeth of-
fer still another great advantage for the student of race. They are
just as firmly controlled genetically as perishable blood groups
and fingerprints. No environmentalist, however biased, can dem-
onstrate that racial peculiarities in dental details are not strictly
hereditary.
Tooth size is not of primary consequence in determining evolu-
tionary grades, but tooth form is of major importance in tracing
racial lines of descent. In fact, so great are the differences in tooth
6 Following is a short bibliography on racial variations in teeth.
M. deTerra: Beitrdge zu einer Odontographie der Menschenrassen (Berlin:
Berliner Verlagsanstalt; 1905).
T. D. Campbell: “Dentition and Palate of the Australian Aboriginal,” PKSF
No. 1 ( 1925).
M. Heilman: “Racial Characters in Human Dentition,” PAPS, Vol. 67, No. 2
(1928), pp. 157-74.
M. R. Drennan: The Dentition of a Bushman Tribe,” ASAM, Vol. 24, Pt. 1
(1929), PP- 61-87.
A. A. Dahlberg: “The Dentition of the American Indian,” in W. S. Laughlin,
ed.: Papers on the Physical Anthropology of the American Indian (New York:
Viking Fund; 1951).
I. Gleiser and E. E. Hunt: “The Permanent Mandibular First Molar, Its Calci-
fication, Eruption, and Decay,” AJPA, Vol. 13, No. 2 (1955), pp. 253-84.
M. Klatsky: “The Incidence of Six Anomalies of the Teeth and Taws ” IIS
Vol. 28 ( 1956), pp. 420-8.
G. W. Lasker and M. M. C. Lee: “Racial Traits in the Human Teeth,” JFS,
Vol. 2, No. 4 ( 1956), pp. 401-19.
C. F. A. Moorrees: The Aleut Dentition (Cambridge, Mass.: Harvard Univer-
sity Press; 1957).
J. C. M. Shaw: The Teeth, the Bony Palate, and the Mandible in Bantu Races
of South Africa (London: John Bale Sons & Danielsson; 1931).
Shaw: “Cusp Development on the Second Lower Molars in Bantu and Bush-
men, AJPA, Vol. 11 (1927), pp. 97—100.
Shaw: ‘Taurodont Teeth in South African Races,” JAnat., Vol. 62 (1028)
pp. 476-96.
I
352 An Introduction to Fossil Man
morphology which first appeared in the earliest fossil specimens
of each area, and which have largely persisted to this day, that
the time of separation of the different lines of descent in the genus
Homo must be set back to a period of which we have no record. I
make this statement because the rates of change in tooth morphol-
ogy are well known from the study of other animal genera. Some
of the differences that I shall describe are both progressive, having
evolved by succession, and racial, having evolved by branching,
whereas others are racial only.
These variations include : ( 1 ) differences in the relative sizes of
individual teeth or groups of teeth (incisors, canines, premolars,
and molars) in the same tooth row of either the upper or the
lower jaw; (2) differences in the length of the cheek teeth (from
the first premolar through the third molar) of the upper jaw in
proportion to the anterior skull length as measured from basion
(the forward lip of the foramen magnum) to nasion; (3) dif-
ferences in the crown patterns of the various teeth; and (4) dif-
ferences in the roots and pulp cavities.
In the Australopithecines, including Zinjanthropus, the molars
and premolars are very large in comparison to the canines and
incisors. This disproportion is not found in any human population,
either of Homo erectus or of Homo sapiens. In this respect human
teeth resemble ape teeth more than the teeth of their fellow Homi-
nids.
Among living Mongoloids the front teeth (incisors and canines)
are even larger, compared to the cheek teeth (premolars and
molars), than in other racial lines. The Australoids vary in the
opposite direction, not because their front teeth are small, but
because their cheek teeth are particularly large. Among Mongol-
oids the third molar is often congenitally lacking in one or more
tooth rows; a fourth molar turns up now and then among Austra-
lian aborigines. Among Caucasoids the upper lateral incisor is
usually much smaller than the upper medial incisor, whereas in
P. O. Pedersen: “The East Greenland Dentition,” MOG, Vol. 142, No. 3
(1949). PP- 1-256.
E. K. Tratman: “A Comparison of the Teeth of People [of] Indo-European
Racial Stock with [Those of] the Mongoloid Racial Stock,” DR, Vol. 70 (1950),
Nos. 2-3, pp. 63-88.
Lines and Subspecies of Fossil Men 353
other races the difference in size between these two teeth is much
less.
The second set of variations is not limited to the teeth, but com-
pares the combined mesiodistal length of the upper cheek teeth to
a sagittal dimension on the skull, the basion-nasion chord. The
formula, length of the premolar = molar row X 100 -f- BN length,
is known as Flower’s index, after its inventor, H. W. Flower, who
was a great zoologist accustomed to classifying mammals in gen-
eral and not limited to the minuscule zoological realm of man.7 He
divided the range of his index into three parts: below and includ-
ing 41-9 per cent is microdont; from 42.0 through 43.9 per cent is
mesodont; and 44.0 per cent and upward is megadont.
TABLE 13
FLOWER’S INDEX
Males Females Males & Females
Polynesians
Non-British Europeans
Ancient Egyptians
British
Central & South Indians
40.1%
40.5
40.8
41.0
41.4
41.6
41.2
41.6
41.3
41.0
41.3
Microdont
Bushmen, South Africa *
42.4
Chinese
42.6
American Indians
42.8
Mesodont
African Negroes
43.2
44.6
43.9
Javanese & Sumatrans
43.3
Melanesians
44.2
Andamanese
44.4
46.5
45.5
Megadont
Australian Aborigines
44.8
46.1
45.5
Tasmanians
47.5
48.7
48.1
* A composite of 29 individuals from four series. Drennan: op. cit.
On the whole the Caucasoids are microdont, the Bushmen and
Mongoloids mesodont, and the Australoids and their Melanesian
and Negrito neighbors and kinsmen megadont. The Polynesians
are classified as microdont because, although their teeth are large,
their basion-nasion chord is very long. The Negroes of Africa
7 H- w- Flower: “On the Size of Teeth as a Character of Race,” JRAI, Vol. 14
(1885), pp. 183-6.
de Terra: op. cit., pp. 183-6.
354
An Introduction to Fossil Man
straddle the mesodont-megadont line, with the males on one side
and the females on the other.
I
Racial Variations in the Form and Structure of Teeth
The morphological differences in the crowns, roots,
and internal structure of human teeth constitute an enormously
complex subject to which a number of specialists have devoted
their lives, and which can only be summarized here, with little
detail. On Table 14 are listed a few notations concerning the
TABLE 14
RACIAL VARIATIONS IN TOOTH FORM
Caucasoid Mongoloid Australoid Negro Capoid
Shoveling rare
extreme
probably
rare*
Ridging
present
common
Premolar Cone
present
Cingulum (Collar)
rare
W rinkling
rare
rare
Enamel Pearl Ainu
present
present
Enamel Extensions
present
Short Roots
present
Extra Roots
present
Taurodontism rare
present
rare
rare
Cusp Formula 4-4-3
4-4-3
4-4-4
4-4-4
4-4-3
5-4-4
5-4-5
5-4-5
5-5-5
5-5-5
Missing Third Molars — 20 %f
-70%
-13%
3% 7
10% ?
Fourth Molars
rare
6 & 7 Cusps 3%
-20%
37%
16% ?
?
Carabelli’s Cusp common
rare
rare
* Common among full-sized Capoid skulls, but not, apparently, among modern Bushmen, whose teeth
are usually too worn to tell.
t Heilman (1928) cited a figure of 49 per cent for a series of ninth century a.d. Hungarian skulls, but
we do not know how many of them were Mongoloid.
variations in fifteen items of dental morphology in the five major
human subspecies postulated in Chapter 1. The term Negro is
used instead of Congoid, because we are speaking of African
Negroes only. For them and for the Capoids (Bushmen) less
abundant and less complete data are available than for the other
three groups. The Caucasoids include Europeans, Near Easterners,
Hindus, and Tamils. In dental morphology these groups are all
Racial Variations in the Form and Structure of Teeth 355
Fig. 49 Racial Variations in Tooth Structure: Shoveling and Ridging.
These drawings represent only upper median incisors. A. Two views of a
Sinanthropus tooth, with raised edges and a basal bulb or tubercle. (After Weiden-
reich, 1937.) B. Two Sinanthropus teeth with incurved edges and basal
tubercles in the form of teats or swollen ridges. (After Weidenreich, 1937.)
C. Four modern Aleut teeth with raised or wrap-around edges, but without basal
tubercles. D. A shoveled incisor, partly worn, seen from the occlusal view and
greatly enlarged. The dentine is cross-hatched. Note that in the shoveling
phenomenon the thickness of the enamel remains constant; the dentine is shoveled
as well as the enamel. (After Tratman, 1950.) E. A modem tooth showing a medial
ridge, commonest among Negroes but also found in other races, including Euro-
peans. (After Weidenreich, 1937.) F. An Upper Paleolithic European tooth from
the so-called Negroid youth of Grimaldi, with double ridging and without raised
edges. This tooth form is also seen in other European skulls, particularly among
Neanderthals. Its presence does not necessarily make the Grimaldi youth Negroid.
alike. The Ainu have been included with the Caucasoids for a few
features. The Mongoloids include eastern Asiatics, Malayans,
Indonesians, Polynesians, Eskimo, and American Indians. As a
whole they are alike, except that among the Polynesians some of
the typical Mongoloid features are attenuated; among some local
American Indian populations the usual Mongoloid features have
beome exaggerated; and among at least two isolated Eskimo
populations local tooth patterns have arisen.
Shoveling is a trait that affects the incisors and canines only,
and the upper teeth, as a rule, more than the lower ones. In a mild
form of shoveling the inside lateral edges of the tooth are bent
356
An Introduction to Fossil Man
backward to form a pair of rims, leaving a concavity between. In
more extreme cases the edges are wrapped around to such an
extent that their borders face inward, and such a tooth looks,
in section, like a Gothic capital C lying prone, C. J . In the
prone-C type the borders can meet at the root end and part com-
pany halfway up, like the flower of a calla lily. Or they can be
fused, and the tooth is tubular or barrel-shaped (see Plate XXIV).
Another extreme form is double-shoveling, found in some south-
western American Indians; among them a concavity on the front
or labial side matches the usual one on the back or lingual side.
For laymen accustomed to seeing the teeth of Caucasoids and
Negroes only, these extreme forms have almost to be seen to be
believed.
Shoveling is often accompanied by from one to three teatlike
protrusions rising from the base of the crown or by vertical ridges
or ribs running from the base to the cutting edge on the lingual
side. As teats and ridges are part of a single complex, they have
been so listed. Among Negroes one often finds them present with-
out shoveling. In South Africa, shoveling turns up in Bantu teeth,
possibly as a result of mixture of Bantu and Bushmen.
Among Asiatic Mongoloids and American Indians shoveling
reaches a level of 90 to 100 per cent. Among many groups of north-
ern Europeans and among native white Americans a simple, un-
spectacular form of shoveling, involving merely a raising of
borders, reaches an incidence of 10 per cent or more, but this
figure may be less than the frequency of the gene or genes that
cause it. In children born in Japan of American fathers and Japa-
nese mothers, the Caucasoid form, being a chisel-like nonshoveled
incisor, appears in the majority.8
What I have called in Table 14 a premolar cone is a rod or
cone or teatlike excrescence of enamel protruding from the center
of the groove in the middle of the occlusal surface of a premolar,
more often mandibular but sometimes maxillary, and usually in
the second premolar. When it is present in the first premolar it is
8 K. Hanihara: “Studies on the Deciduous Dentition of the Japanese and
Japanese-American Hybrids,” ZZ, Vol. 63 (1954), pp. 168-85; Vol. 64 (1955),
pp. 63-82, 95-116; Vol. 65 (1956), pp. 67-87; Vol. 65 [sic] (1957), pp. 151-64.
After G. W. Lasker: “Recent Advances in Physical Anthropology,” BRA, 1959,
pp. 1-36.
Racial Variations in the Form and Structure of Teeth 357
Fig. 50 Racial Variations in Tooth Structure: the Premolar Cone.
A. A first premolar of an American Indian, in section and greatly enlarged. B. The
same tooth, side view. C. The same tooth, occlusal view. D. An Aleut premolar
with premolar cone. The premolar cone is a Mongoloid peculiarity. A cone
rising from the center of an upper first premolar, it is visible in young teeth only, as
it is soon broken off or worn down. As A shows, it consists of both dentine and
enamel. It is disadvantageous because its destruction leads to a premature exposure
of dentine. (Drawings A, B, C after Tratman, 1950; D after Moorrees, 1957.)
also to be seen in the second. Tratman ( 1950) called it a “dilated
composite odontome,” too technical a term for this book; and
Pedersen (1949) labelled it an “enamel pearl,” not to be recom-
mended because the term is already in use to describe another
structure. In any case, the premolar cone, owing to its position, is
soon worn down, and can be detected with any conviction only
in newly erupted teeth. Like several other bizarre dental details,
it is confined to the Mongoloid subspecies.
A cingulum is a collarlike rim about the base of the crown of
any tooth except the incisors. It may be partial or complete. It
again is a Mongoloid peculiarity, although it may be expected
in some very ancient teeth of other lines. Wrinkling is another
Mongoloid phenomenon, commoner in ancient than in modern
specimens. In some of the Mongoloid molars the surface, instead
of being smooth and flat, is broken up into fine relief by a pattern
of wrinkles. This is also a characteristic of both Australopithecine
and orangutan teeth and may be considered an ancient feature.
An enamel pearl is a pearl-like excrescence on the labial side of
a molar tooth (see Plate XXIV). It is common among Ainu, oc-
curring with a frequency of 30 per cent, and among Eskimo. It
358
An Introduction to Fossil Man
ENAMEL EXTENSIONS
Fig. 51 Racial Variations in Tooth Structure: the Cingulum, Wrinkling,
Taurodontism, and Enamel Extensions. A. Cingulum in an upper canine of
Sinanthropus. B. Cingulum in a lower molar of Sinanthropus. C. Wrinkling in the
crown of a third molar of Sinanthropus. D-H. Molar pulp cavities in a series of
teeth from cynodont to taurodont. I. An X-ray of a taurodont tooth. J-K. Enamel
extensions on the roots of molar teeth, Sinanthropus. A cingulum is an enamel
collar at the base of the crown of a tooth. It may run all the way around or, more
frequently in man, only part way. Wrinkling, like shoveling and the premolar
cone, involves both enamel and dentine. Teeth which have this feature are
wrinkled when cut, and the wrinkles soon wear down. Its function is unknown,
but it is found in the molars, and sometimes other teeth, of apes, Australopithecines,
and men. It is commonest among Mongoloids. Taurodontism is a neotenous condi-
tion of the molars and sometimes the premolars. The pulp cavities of taurodont
teeth become enlarged because the roots fail to grow to fully adult height; the
underside of the root area fails to develop completely. Thus the pulp cavity of the
tooth is elongated downward from above. As the crowns wear, the dentine
hardens, and the tooth can be worn down to the gums and even onto the roots.
D is the molar of a European with normal pulp cavities. Such a tooth is called
cynodont (dog-toothed). E is the molar of a Bushman-Bantu hybrid, with slight
taurodontism ( bull-toothedness ) . F is a tooth from the Heidelberg jaw, with
medium taurodontism; and G and H are teeth from the jaw of a teen-aged
Krapina youth, with extreme taurodontism. His molars had erupted at an early and
incomplete stage of root development. I is the drawing of an X-ray of a taurodont
tooth. Enamel extensions, found among Mongoloid teeth particularly, are extensions
of the enamel of the crown onto the roots. In some cases the enamel goes under
the gap between the roots like a cinch strap; in others it extends down the outside
Racial Variations in the Form and Structure of Teeth 359
has also been found in other Mongoloids and among Bushmen.
Among Mongoloids the lower border of the enamel-covered por-
tion of the molar crown dips down, in many cases, onto the neck
of the tooth and onto its root. In other races the lower border of
the enamel forms a straight line. When the border of the enamel
dips down, it forms what is called an enamel extension.
The roots of Mongoloid teeth also tend to be short for the size
of their crown; this is also characteristic of Lapps in Europe. Mon-
goloid molars also tend to have one or more extra roots in the for-
ward part of the molars. They also lead the others in the fre-
quency with which upper third molars are congenitally missing —
up to 70 per cent in some populations — whereas this progressive
trait is much rarer in all the others. The opposite tendency, to have
four molars on any one side of either jaw, is rare everywhere but
commonest among Australian aborigines.
In the evolution of the horse and other grazing animals teeth
with long crowns, or rather high ones, which would give more
years of wear than those with short or low crowns, became prev-
alent through the usual process of natural selection. In human
beings, for whom the wear on the molars and premolars is critical,
the tooth had to grow larger or its crown had to become higher. In
the Australopithecines, particularly in Kromdraai and Zinjanthro-
pus, the molars and premolars simply became huge. The genus
Homo took the alternative solution, known as taurodontism, or
bull-toothedness. Ordinary molars are cynodont (dog-toothed).
As Gleiser and Hunt ( 1955) discovered, taurodontism is an in-
fantile character, a delay in the growth rhythm of the roots and
inner crown structure of a tooth. In the forming molar or pre-
molar, before it has erupted, the occlusal surface of the crown has
already assumed its adult form but the pulp cavity is much larger
than it will be later, under normal circumstances of growth, and
the roots have failed to separate and to become defined on their
inner surfaces. The roots are more or less barrel-shaped. If this in-
fantile form is retained into later life, the large pulp cavity reach-
of the root to its tip. Enamel extentions give the tooth additional wear, par-
ticularly when chewed on the side. The most notable examples of these are
among Sinanthropus teeth and those of the East Greenland Eskimo. (Drawings
A, B, C, J-K after Weidenreich, 1937; D-H after J. M. Shaw, 1927, and Tratman
1950; I after Tratman, 1950. )
An Introduction to Fossil Man
360
ing well down into the combined or undifferentiated root may
become an asset. As the enamel wears off, the pulp hardens
to dentine, and the wear may be carried well past the point of
danger in an ordinary tooth.
As one would expect, taurodont teeth are characteristic of
Mongoloids, particularly of such hard-chewing Mongoloids as
Eskimos. They are infrequent in living Caucasoids, but we shall
encounter them in certain hard-chewing dead ones. Among Ne-
groes they seem to occur only among southern Bantus, who are
part Bushman anyway, whereas among the Capoids they are
characteristic only of full-sized ancestors of the Bushmen un-
earthed by archaeologists.
In contrast to the dental details that we have so far summarized,
the pattern taken by the cusps on the crowns of the upper and
lower molars, and the numbers of these cusps on each tooth, have
attracted by far the most attention from tooth experts concerned
with human evolution and racial variation. The bibliography is
legion, because it includes the efforts of paleontologists, zoologists,
and dentists as well as of anthropologists.
As Gregory long ago demonstrated, the primordial crown pat-
tern of human lower molars is a set of three grooves in the form of
a Y lying on its side, with its tail pointing forward and its two arms
pointing to the rear. In each obtuse angle are stationed two cusps,
and in the acute angle a fifth cusp. This pattern, known as Y-5, is
also called the Dryopithecus pattern because it was characteristic
of that family of Hominoids from which we may all have been
descended — the Dryopithecinae ( see Chapter 6 ) .
In the Australopithecines this pattern was altered by the addi-
tion of extra cusps and grooves, apparently to compensate for the
added burden of chewing and grinding imposed by the coarse
diet necessary for life on the ground. In the earliest specimens of
Homo yet found, which we shall study in the next four chapters,
no increase in molar crown complexity comparable to that of the
Australopithecines can be observed. Some teeth show the opposite
tendency, a reduction in complexity of the original Dryopithecus
pattern. In all modern races, reduction has proceeded in nearly all
the molar teeth, in varying degrees.
In modern teeth the molar crown patterns have been simpli-
Racial Variations in the Form and Structure of Teeth 361
Fig. 52 Lower Molar Crown Patterns. Four stages of development of lower
molar teeth, illustrating the different changes from the Dryopithecus to the most
advanced pattern. (A) Dryopithecus pattern indicated by Y5, meaning primitive
system of grooves and cusp formula. ( B ) Modified Dryopithecus pattern indicated
by Y4, meaning cusp formula reduced to four and groove system primitive with the
loss of the posterior limb of the Y. ( C ) Primitive cusp formula retained but
groove system changed to cruciform, the sign being +5. (D) Cusp formula and
groove system changed, sign +4. (f.a.) fovea anterior; (f.p.) fovea posterior.
(Drawings and captions from Milo Heilman: “Racial Characters in Human Denti-
tion, PAPS, Vol. 62, No. 2 ( 1928), p. 165. Courtesy of the American Philosophical
Society. )
fied in two ways: (1) the groove pattern has changed from a lazy
Y (to use a Western cattle-branding term) to a simple cross, +;
( 2 ) the number of cusps has dropped from five to four, or even to
three or two. With five or four cusps and Y’s and crosses, four com-
binations are possible: Y-5, Y-4, +5, and +4. Because of the sym-
bol used, cross-5 and cross-4 are usually referred to as “plus five”
and plus four, which is confusing because plus implies num-
ber rather than form.
In every race for which pertinent data are available, crown-
pattern reduction is greater in the upper than in the lower jaw,
and only in the lower molars have both the groove patterns and
the cusps numbers been systematically examined. In every race
studied, the first lower molar has the original, Dryopithecus, Y-5
pattern in the majority of instances. In Caucasoids its frequency
is about 7 5 per cent, and in all other races it is between 95 and
100 per cent. This is our most conservative molar tooth.
Y-4 is a rare pattern. In Europeans it appears in about 12 per
cent of the first molars, and in African Negroes in the same per-
362
An Introduction to Fossil Man
centage of the second molars. Otherwise, its appearance is spo-
radic. From the standpoint of reducing the mesiodistal length of
a molar crown, Y-4 is less efficient than the cross pattern, and
both +5 and +4 greatly outnumber it in all races.
In all races studied, +4 outnumbers +5 in the second molars.
However, in the third molars the +5 pattern outnumbers the +4
pattern in all populations studied except the Europeans, Chinese,
and some Eskimo groups. Considering the three lower molars as a
whole, only among the Caucasoids do we find more teeth with
four cusps than with five, and more groove patterns in the form of
a cross than with Y patterns. In general, the lower molar crowns
are most reduced in the Caucasoids and least reduced in African
Negroes and Australian aborigines. Among the Bushmen, whose
teeth are very small, we find the highest ratio of five-cusped lower
molars. Unfortunately, we do not know the frequencies of their
groove patterns.
In the upper molars all races have a majority of four-cusped
crowns, nearly 100 per cent, in the first molars. In the second
molars, however, racial differences in the number of cusps appear;
among Caucasoids and Mongoloids three cusps are seen in 30 to
42 per cent of these teeth, whereas in Negroes the ratio decreases
to 22 per cent and in Australian aborigines and Bushmen all up-
per second molars are apparently four-cusped. In the third mo-
lars more than half the teeth have three cusps among Caucasoids,
Mongoloids, and Bushmen, whereas among Negroes and Aus-
tralian aborigines more than half have four cusps.
From this statistical exercise we see that reduction in the crown
pattern is most advanced among Caucasoids, least advanced
among Negroes and Australian aborigines, and intermediate
among Mongoloids. However, among Mongoloids and Bushmen
the third molar is much more reduced in comparison to the first
and second molars than in the other races, including the Cauca-
soid.
From the over-all cusp formula given on Table 14, it is ap-
parent that the Caucasoid lower molar crowns are the most re-
duced of all, and those of the Negroes the least reduced. Among
the Mongoloids and Bushmen the reduction is concentrated in the
upper third molar.
Racial Variations in the Form and Structure of Teeth 363
In all races there is a tendency for the upper third molar to be
congenitally absent, but this tendency is most marked in the Mon-
goloids, particularly the Eskimos. Fourth molars are rare in all
races, but are found most often among Australian aborigines.
They too lead in the number of teeth having six or seven cusps,
which is one or two more than the standard number for the
Dryopithecus pattern.
Among Caucasoids, and only sporadically in other races, a
special feature known as Carabelli’s cusp ( described in Chapter
7; see Plate XXIV), is seen; among Europeans its frequency is as
high as 40 per cent. Carabelli s cusp is an accessory cusp situated
on the front part of the inside or tongue-surface of upper molars.
Carabelli s cusp is situated below the level of the surface of the
crown when the tooth is freshly erupted and unworn. When the
tooth has been worn down to its level, this accessory flange serves
to widen the occlusal surface of the molar and thus slow down
further wear. In this way it performs the same function as tauro-
dontism in prolonging the useful life of the tooth. It is commonest
on upper first molars and rarest on upper third molars. It is
only found on upper second molars if it is also present on upper
first molars; and when it turns up on upper third molars it is also
to be seen on the other two molars of the row.
In summarizing this survey of racial tooth morphology, we are
struck with several facts. The teeth of Caucasoids are plain and
simple, with a single peculiarity, Carabelli’s cusp. Of all races the
Caucasoids show the greatest over-all reduction in tooth-pattern
details, although their teeth are not the smallest in the world.
Negro and Australian aboriginal teeth are also relatively simple,
although less reduced than those of Caucasoids; indeed Negro
teeth seem to be most like those of primitive Caucasoids.
Mongoloid teeth, however, are far from plain. They are deco-
rated with a myriad of complicated details. They are quite dif-
ferent from those of the three races just mentioned, with several
vaiiations and exti ernes of shoveling; with premolar cones; wrin-
kling of molar enamel; enamel extensions onto the roots; enamel
pearls; short, taurodont roots; and a tendency for the upper third
molars to be greatly reduced in crown pattern as well as in size, or
even to be congenitally absent. Mongoloid teeth are not so much
364
An Introduction to Fossil Man
primitive as they are deviant from those of the rest of Homo
sapiens. Bushmen teeth, although very small, resemble those of
Mongoloids morphologically more than they do the teeth of Cau-
casoids, Negroids, or Australoids.
Were one to classify human races on the basis of teeth alone, it
would be easy to place the Mongoloids and Capoids in one cate-
gory and all the other races in another. Differences in the incisors
and canines alone, without reference to the cheek teeth, widely
separated the Mongoloids from the Caucasoids, and as we shall
presently show, these racial differences in tooth form go back as
far as we can trace the ancestors of man.
1 1
Facial Flatness as a Criterion of Race
A general peculiarity of Mongoloid dentition is that the ac-
cent is on the front part of the mouth. The incisors and canines
are large and elaborately braced for hard work, whereas the mo-
lars taper off rapidly from the first to the third, and the latter is
often missing. Along with this forward orientation of the teeth
goes a forward stance of the temporal and masseter muscles which
furnish most of their motor power. In the forward part of their
area of attachment on the frontal bone the temporals invade the
forehead on each side, above the outer halves of the orbits, thus
making the forehead look narrow, and incidentally pressing the
outer portions of the orbits from behind and making them rela-
tively shallow. As part of the same complex, the masseters are
hung from forward-jutting zygomatic bones, the well-known
“high cheek-bones” of the Mongoloids.
All these specialties, in combination with a low-bridged nasal
skeleton, give the Mongoloids, and the Bushmen as well, an ap-
pearance of facial flatness superficially reminiscent of the apes
and Australopithecines. In the case of the Australopithecines this
resemblance is superficial because those hominids had very large
cheek teeth and small front teeth. At the other extreme, Cauca-
soids have pointed, sometimes beaklike faces which remind one
of some of the lower primates, particularly the South American
monkeys, and of Proconsul.
Facial Flatness as a Criterion of Race 365
Apart from the teeth, the degree of facial flatness is probably
the best criterion of race which the skull offers for study because
it seems to have little to do with evolutionary grades. With this in
mind, T. L. Woo and G. M. Morant published, in 1934, the results
of their monumental study of facial flatness in 131 series of skulls,
totaling 5,788 specimens, and covering all major racial groups.9 In
order to describe this racial criterion accurately, they devised
four indices, each based on the measurement of a chord between
matched points on either side of the facial skeleton and the sub-
tense drawn between the center of this chord and a given point
on the sagittal line of the facial skeleton in front of that chord.
The formula for each index is: subtense X 100 chord. The four
chords and their subtenses are as follows.
( 1 ) The internal biorbital chord, that is, the distance between
the points on the outer edges of the orbits where the frontal and
malar bones meet. The subtense used with this chord is that
formed by bisecting the triangle created by joining these two
points to nasion.
(2) The simotic chord, that is, the minimum horizontal
breadth of the paired nasal bones. The point used for the sub-
tense is the spot on the nasal suture lying exactly between and in
front of them.
(3) The midorbital chord, that is, the distance between the
points on the lower border of the orbit where malar and maxilla
meet. The subtense is to the lower tip of the suture between the
nasal bones.
( 4 ) The “ face breadth ” of Martin, that is, the chord between
the points on the zygomatic-maxillary suture which are lowest in
reference to the eye-ear plane. The subtense is to alveon, the point
on the outer sagittal edge of the maxilla between the upper lateral
incisors.
The indices derived from these four sets of chords and sub-
tenses are called: (1) frontal index of facial flatness, (2) simotic
index, (3) rhinial index of facial flatness, and (4) premaxillary
index of facial flatness. Table 15 shows the numbers of series and
ranges of means for Caucasoids, Mongoloids, Australoids and
n,' ip ,W°° and G' „M- Morant: “A Biometric Study of the ‘Flatness’ of the
Facial Skeleton m Man, Biometrika, Vol. 26 (1934), pp. 196-250.
366
An Introduction to Fossil Man
Fig. 53 The Four Indices of Facial Flatness. Above: The chords (left) and
subtenses ( right ) from which the four indices of facial flatness are calculated. The
order is from top to bottom: No. 1 is the frontal index, No. 2 the simotic, No. 3 the
rhinial, and No. 4 the premaxillary. Below: The chords and subtenses laid out.
Number 2, because of its small size, is scaled three times the size of the others.
The skull is that of a Vedda of Ceylon. Its indices of facial flatness are: (1) 22;
(2) 38; (3) 30; and (4) 34. These figures identify it as Caucasoid.
Oceanic Negroids, African Negroes and Capoids. In the Cauca-
soid segment the Ainu and the ancient Egyptians have been listed
separately because they show a number of differences from the
others. The combination of India and Ceylon ( including Veddas )
has also been listed separately, mainly to indicate that their
inhabitants are essentially the same in these indices as the Euro-
peans and Near Easterners. In the Mongoloid segment the Nepa-
lese are given separate status because they are of mixed Cauca-
soid and Mongoloid ancestry. In the same way, an ancient Nu-
bian series has been separated from the Negroes because the
Nubians were partly Caucasoid.
367
Facial Flatness as a Criterion of Race
TABLE 15
THE FOUR INDICES OF FACIAL FLATNESS
OF WOO AND MORANT
No- n #2 #3 H
Ser. Frontal I Simotic I Rhinial I Premax. I
Ainu
1
17.2
36.6
32.2
34.0
Ancient Egyptians
6
17.4-18.3
32.1-41.1
30.3-40.3
35.8-37.9
India and Ceylon
9
19.6-21.0
37.5-49.1
35.4-43.5
35.3-38 4
Europe and West Asia
32
17.4-20.5
40.0-53.0
37.8-48.6
33.4-37.2
Asiatic, Mongoloid and
Eskimo
23
14.1-16.5
25.5-34.6
25.4-33.3
31.4-35.8
Nepalese
1
16.9
37.7
31 5
3.3 ]
American Indians
6
16.7-17.6
29.5-49.0
32.4-39.4
34.3-36.0
Polynesians
11
16.4-18.0
36.7-43.4
31.5-39.4
34.3-37.0
Australians and Tas-
manians
Papuans and Mela-
7
17.6-18.7
35.5-44.4
29.9-37.2
38.7-42.3
nesians
Negritos, Philippines
9
17.2-18.4
26.1-39.8
31.1-37.4
35.7-37.9
and Andamans
4
15.8-17.2
23.9-35.5
28.1-34.2
34.1-34.7
Nubians
2
17.7-18.3
30.2-36.3
33.6-34.6
34.5-35.3
Negroes
14
17.0-18.1
18.6-33.6
24.2-30.8
34.1-39.1
Bushmen and Hotten-
tots
4
15.5-17.0
16.9-29.3
19.2-27.9
34.1-35.8
This table can be most easily understood in terms of the four
charts on page 368. They show clearly that, in the first three
indices, the Caucasoids occupy an extreme position with by far
the most pointed, or least flat, faces of all. In the first three indices
the Caucasoid means do not even overlap those of the Bushmen-
Hottentots and Asiatic Mongoloids and Eskimos. Only in the
fourth index do the Caucasoids occupy an intermediate position,
because this index is influenced by prognathism and thus tends to
reflect evolutionary grade more than the other three do. In this
index, as in so many other criteria, the Mongoloids and Bushmen
stand at one extreme; but instead of the Caucasoids it is the Aus-
tralian aborigines and Tasmanians who stand at the other. The
Caucasoids and Polynesians are located in the middle of the scale.
In summarizing the information yielded by these four indices
of facial flatness, we find that some races are far more deviant
from a standard, generalized Homo sapiens face form than are
WOO'S AND MORANT’S FOUR INDICES OF FACIAL FLATNESS
1. FRONTAL INDEX OF FACIAL FLATNESS
Ainu, Caucasoids ■
Australians, Tasmanians ■■
Papuans, Melanesians ■■■■
Negroes ■§^■■1
Polynesians
American Indians
Negritos, Philippinos, Andamans
Bushmen, Hottentots (■■■■■■■■■■■
Asiatic Mongoloids, B
Eskimo, Nepalese
2. SIMOTIC INDEX
Ancient Egyptians, Ainu, Caucasoids ■
Australians, Tasmanians
Polynesians
American Indians
Papuans, Melanesians
Philippine Negritos, Andamans
Asiatic Mongoloids, Eskimo, Nepalese
Negroes, Nubians | I
Bushmen, Hottentots
3. RHINIAL INDEX OF FACIAL FLATNESS
Egyptians, Ainu, Caucasoids ■ |
American Indians
Polynesians ■■
Papuans, Melanesians HH
Australians, Tasmanians
Negritos, Philippinos, Andamans ■■■■■■
Asiatic Mongoloids, Eskimo
Negroes, Nubians
Bushmen, Hottentots HHHIHHHHHHBHi
4. PREMAXILLARY INDEX OF FACIAL FLATNESES
Australians, Tasmanians
Negroes
Papuans, Melanesians
Caucasoids
Polynesians
American Indians ■■■■■
Bushmen, Hottentots ■■■■■■■I
Asiatic Mongoloids, Eskimo
Negritos, Philippinos, Andamans ■■§
Racial Origins and Racial Continuities 369
others. The ranking, from most to least deviant, is: Caucasoids,
Australoids, Capoids, Asiatic Mongoloids and Eskimos, Asiatic
Negritos, African Negroes, American Indians, Papuo-Melanesians,
and Polynesians. If we eliminate the fourth index on the ground
that it confuses grade with line, the Caucasoids appear even more
distinctive.
This ranking suggests that Caucasoids, Australoids, and a Mon-
goloid-Capoid combination are extreme or primary forms of man-
kind, whereas Ainu, Polynesians, Papuo-Melanesians, African Ne-
groes, Asiatic Negritos, and even American Indians are inter-
mediate forms. The extreme differentiation of the Caucasoids in
degree of facial flatness matches that of the Mongoloids in dental
anatomy, and is equally ancient.
Racial Origins and Racial Continuities
In this chapter a framework has been erected for the de-
tailed study of the origins and evolutionary progress of each of
the five geographical races of man, as defined in Chapter 1. We
have shown that from the start of the Middle Pleistocene the Old
World, in which man arose, was divided into five breeding
grounds, sufficiently separated from one another by physical and
climatic barriers to permit a human subspecies to evolve, almost
but not quite independently of its neighbors. We have traced the
archaeological sequences in each region, and shown that in them
tool-making techniques followed essentially but not wholly in-
dependent paths.
Short-cutting anthropometric details, we have set the boundary
between Homo erectus and Homo sapiens on the basis of brain
size, the degrees of curvature of the bones composing the cranial
vault, and, to a lesser extent, on tooth size, particularly as tooth
size is related to brain size. We have explored and rejected the
concept that Homo sapiens must, by definition, have had a chin,
and have shown how chins came into being after the races of man
had crossed the erectus-sapiens threshold.
From the consideration of evolutionary grades we have moved
on to lines of descent, and have selected two out of many criteria
of racial differentiation that are particularly easy to follow. These
370
An Introduction to Fossil Man
are tooth morphology and degrees of facial flatness. In all regions
where they can be traced, both persist from the first appearance
of man to the end of the Pleistocene.
With this lengthy introduction we have bridged another gap
between the study of our prehuman ancestors and the details of
human history. From here on we shall pursue the second subject,
not by grades, for that framework has been sufficiently explored,
but by racial lines.
Pygmies: Onges from Little Andaman Island.
XVIII
A Pygmy from the Congo.
XX
>11
X and 6
II )l II It (III
7 8 9 10 ,1 ]2
Mil ki
13 14 15
I! II il
16 17 18
II It «» u |
19 20 2, 22 y
The Karyotype of Man
(American Negro, Male).
The forty-six chromosomes
of the human cell nucleus,
shown as they appear under
the microscope and arranged
by pairs in order of size, ex-
cept for the sex chromosomes
X and Y, which are of differ-
ent sizes. A female would
have two X chromosomes.
Research in differences in de-
tails of chromosome struc-
ture in man has not yet pro-
gressed to the point at which
we can identify racial differ-
ences.
Human chromosomes.
XXI
Zinjanthropus palate, compared with that of an Australian aborigine.
XXIII
a. Extreme shoveling of the
upper incisors of a Pima In-
dian whose lateral incisors
are barrel-shaped.
b. In the teeth of another
Pima Indian shown here, the
upper median incisors are
double-shoveled — they are
concave on both the outer
and the inner surfaces.
c. Carabelli’s cusp, on the upper first molar of a white
American.
d. e. f. The enamel pearl, a globular form of enamel ex-
tension, on the molars of East Greenland Eskimos.
SOME RACIAL PECULIARITIES IN TOOTH STRUCTURE
XXIV
v mmrn
Skull of La Fi
c. Upper Cave Male
d. Skhul 5
FLESH RECONSTRUCTIONS OF FOSSIL MEN
By Maurice Putnam Coon
a. Steinheim
b. Wadjak 1
b. Circeo 1
d. Cro-Magnon
c. Combe Capelle
a. La Chapelle aux Saints
FLESH RECONSTRUCTIONS OF FOSSIL MEN
By Maurice Putnam Coon
XXXI
The Alpha and Omega of
Homo sapiens: An Australian
aboriginal woman with a
cranial capacity of under 1,-
ooo cc. (Topsy, a Tiwi) ; and
a Chinese sage with a brain
nearly twice that size (Dr. Li
Chi, the renowned archaeolo-
gist and director of Academia
Sinica).
a
&
a
9
K
3g
PITHECANTHROPUS
AND THE AUSTRALOIDS
T/ie Pithecanthropus Line
f the five lines of human descent that we shall now
trace, the best one to begin with is the Australoid because it is the
oldest known, the simplest to follow, and the most archaic today.
The Australoid is the one human subspecies known to be native
to the Oriental faunal region, and it was the first to occupy its
present home, the Australian faunal region.
Owing to extensive post-Pleistocene migrations, the Oriental re-
gion is a racial mosaic, the most complex in the world. West of the
mountains that separate India from Burma fives a medley of peo-
ples, mostly Caucasoids on the plains, with enclaves of Negritos,
Australoids, primitive Caucasoids, and Mongoloids in the hills.
East of the mountains the mainland population is almost solidly
Mongoloid, and so is that of the islands of Indonesia (speaking in
the geographical rather than the political sense ) lying north and
west of Wallace’s Line.
Like India, southeast Asia and the larger Indonesian islands
also contain relict populations tucked away in forest refuges.
These refugee groups are also racially varied, being Negrito, Aus-
traloid, and primitive Mongoloid. As one crosses Wallace’s Line
and moves eastward through the Lesser Sundas toward New
Guinea, the islanders look less and less Mongoloid and more and
more Papuan or Melanesian. New Guinea, Melanesia, and Austra-
lia are (or were) inhabited by a congeries of three principal
kinds of people, Negritos, full-sized Negroids, and Australoids, all
shading into one another in such a way that the Negritos and Ne-
J A C 1 F I C
374
Pithecanthropus and the Australoids
groids form a broken ring around the Australian aborigines.
The Mongoloids of southeast Asia and the islands began to filter
in from the north toward the end of the Pleistocene. We do not
know when the Caucasoids began to occupy their areas of the In-
dian peninsula, but for present purposes this question is unim-
portant, because the Caucasoids did not penetrate east of the
Burmese mountain barrier into the Australoid homeland. All we
are concerned with here are the three patently indigenous
groups: Negritos, Oceanic Negroids, and Australoids.
These three kinds of people differ principally in two features
only, body size and hair form. Variations in body size can be ex-
plained by the mechanisms of dwarfing explained in Chapter 3.
Differences in hair form in this region follow a distinct geographi-
cal pattern in which the Negroid hair is marginal, and therefore
apparently older, than the straight or wavy hair.
It seems reasonable to suppose that these three kinds of native
peoples evolved locally from a common ancestor. Otherwise we
must postulate that several ancestors entered the area in a series
of invasions, each race fully evolved, and carefully avoided one
another until all had reached their present homes. The first ex-
planation requires a certain amount of local evolution and genetic
differentiation, whereas the second demands more migrations
than the archaeological evidence warrants. The second also begs
the question of origins: every race had to originate somewhere.
The solution to this problem lies in the fossil record, which we
shall now pursue, and which tells us nothing about hair form and
little about dwarfs, until the very end. Nevertheless, this fossil rec-
ord, although far from perfect, follows what seems to have been a
single polytypic line through several evolutionary grades, includ-
ing the transition from Homo erectus to Homo sapiens. What we
learn in this chapter can help us in the larger and more compli-
cated areas of eastern Asia, America, Africa, and Caucasoid
Eurasia.
The Pithecanthropus-Australoid Skeletal Material
As Table 16 indicates, thirty-three sites have yielded the
bones of more than a hundred individuals, starting with Pithecan-
375
Pithecanthropus 4
thropus 4, a baby’s skull, and two broken mandibles, and covering
the entire time span from before the base of the Middle Pleisto-
cene to almost the present. Only the sites in Australia containing
patently modern burials, and two dubious sites in Indochina which
contained, respectively, a tooth and a scrap of temporal bone,
have been omitted.
Fossil Men from the Djetis Beds of Java
The oldest fossil-man remains from the Oriental faunal re-
gion, and probably from the world, have been found in Java,
which, among other islands of western and northern Indonesia,
received invasions of Pleistocene animals from both India and
China. The earliest of its hominid-bearing beds is the Djetis,
which contains fossil species believed to have come from the tropi-
cal region of south China, including the pieces of mandible
known as Meganthropus paleojavanicus described in Chapter 7.
The human material found in these beds was: one specimen con-
sisting of two parts of a cranium and two loose incisors, known,
among other names, as Pithecanthropus 4 ( or P-4 ) ; two fragmen-
tary mandibles known as Pithecanthropus B and Sangiran, re-
spectively; and the skullcap of an infant, called Homo modjoker-
tensis.1
Pithecanthropus 4
The larger of the two pieces of Pithecanthropus 4 is the
rear portion of a thick-walled, low-vaulted skull which had been
broken shortly before its discovery, and parts of which had been
lost. The occipital bone is nearly intact, as well as most of the tem-
porals and parietals. What is missing is the frontal and the whole
1 F. Weidenreich: “Giant Early Man from Java and South China,” APAM,
Vol. 40, Part 1 (1945).
G. H. R. von Koenigswald: “Neue Pithecanthropus Funde, 1936-38,” WMDM,
No. 28 ( 1940).
Von Koenigswald: “L’Hominization de l’Appareil Masticateur et les Modifica-
tions du Regime Alimentaire,” in A. Delmas, ed.: Les Processus de I’Hominiza-
tion (Paris: Centre Nationale de la Recherche Scientifique; 1958), pp. 60-78.
Von Koenigswald: “Fossil Hominids from the Lower Pleistocene of Java,”
IGC, No. 9 (London, 1948), pp. 51-69.
Von Koenigswald: Meeting Prehistoric Man (New York: Harper and Brothers;
1956).
PITHECANTHROPUS— AUSTRALOID
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New Guinea Aitape Probably Late Pleistocene 1 ealva H sapiens (?)
Java Wadjak Late Pleistocene or Early No. 1 cranium H. wadjakensis
Recent No. 2 fragments H. wadjakensis
cranium, mandible
378 Pithecanthropus and the Australoids
upper part of the face down to the floor of the nose. The smaller
piece consists of the maxillae, including the palate, the nasal floor,
and ten of the upper teeth, excluding all four incisors and the last
two left molars. Several loose teeth were found in the same site
and apparently fragments of the upper jaws of two other indi-
viduals.
Although there was no point of contact between the two pieces
of P-4, Weidenreich restored the skull as a whole, on the model of
the Sinanthropus skulls with which he had previously worked. He
estimated the cranial capacity at about 900 cc., which matches
the other and later skulls of the same type. The sagittal line of the
Fig. 54 Transverse Section of the Skulls of a Female Gorilla, Pithe-
canthropus 4, and Solo 11. The skull of a female gorilla (inner line) resembles
that of Pithecanthropus 4 in general, except that the latter is more angular than
the former and its sides are more nearly parallel. The largest Solo skull, No. 11
(outside line), is much the same in outline as Pithecanthropus 4, except that it is
more rounded. (Drawing after Weidenreich, 1951.)
skull was keeled, but not crested as in some apes, Swartkrans, and
Zinjanthropus. In fact, the temporal muscles did not extend very
high on the parietals. The occipital torus was pronounced, indi-
cating strong neck muscles, and the brow ridges must have been
protuberant also. The proposed maximum length of 199 mm. and
maximum breadth of 156 to 158 mm. can be matched today in
large-headed Europeans, but the two sets of dimensions are not
comparable. In the Europeans the figures reflect to a large extent
the size of the brain, whereas in P-4 they merely bound the bony
structure of its base. On the sides, the maximum diameter goes
through the crest over the mastoids. The breadth of the brain case,
Pithecanthropus 4 379
in so far as it can be measured, is only about 125 mm. With a
basion-bregma height of only 102 mm. and an auricular head-
height of only 90 mm., it falls within the height range of the larger
Australopithecines, whose heads, being smaller, were relatively
higher vaulted than that of P-4.
The skull in general is formed of a series of planes more or less
rounded at their points of juncture, so that it looks somewhat like
a poorly raised loaf of bread. This is the so-called “ill-filled” look
for which modern Australian aboriginal skulls are also noted. The
base of the skull was relatively flat, and the positions of the fora-
Fki. 55 The Faces of Homo erectus. Only one Homo erectus specimen has a
complete face— Broken Hill (C)— and it lacks a lower jaw. Weidenreich recon-
structed the Pithecanthropus 4 skull, which had a maxilla and palate, with the
addition of the Pithecanthropus B mandibular fragment (A). Using various frag-
ments of face bone and the Sinanthropus 3 skull, he also reconstructed a female
Sinanthropus skull (B). We have no fossil skulls for the erectus stage of either
Caucasoids or Capoids. As shown here, the Pithecanthropus 4 skull is the most
brutal and primitive of the three. The face of the Sinanthropus female is less so,
but partly because of her sex. The Rhodesian skull has the largest brow ridges and
the longest face of all, and its vault is about as high as that of Sinanthropus. How-
ever, it differs from the others in the flatness of the sides of its face. Its zygomatic
arches are weakly developed, not because of small jaw muscles, but because of a
deeply excavated postorbital constriction which gives the temporals room to con-
tract.
men magnum and its flanking occipital condyles indicate, in so
far as they can be trusted to do so, a fully erect posture, although
the nuchal crest, to which the neck muscles are attached, is situ-
ated higher up than in some Australopithecines, notably Zin-
janthropus. Unlike some other fossil skulls and like most modem
380 Pithecanthropus and the Australoids
human races, P-4 has large, downward-pointing mastoid proc-
esses.
The maxillary fragment is notable for its excessive size and mas-
siveness, indicating that the face as a whole must have been large
in all dimensions, with extreme alveolar prognathism. The floor of
the nasal passages was extremely wide ( 36 mm. ) but fully human
in form. In front stands a prominent nasal spine, and the borders
of the opening are guttered, as in living Australoids and Negroes.
The palate is the longest on record, but not the widest. The den-
tal arcade is parabolic, but not as smoothly rounded as in many
other fossil specimens, and the molar rows are nearly parallel.
The most notable feature of this palate is that on each side a
gap, or diastema, separated the canine from the lateral incisor by
5 or 6 mm. Such a gap is normal but not universal in apes, and
even in the gorilla the space is usually narrower than in P-4. It was
found in one of twelve Oreopithecus specimens; it is absent in all
Australopithecine palates; and it turns up rarely in living men:
one such gap has been found in a living Australian aborigine.2
Two other maxillary fragments in von Koenigswald’s possession
are said to be divided in this respect; one has a diastema, the
other lacks it.3 In P-4 this gap was unaccompanied by a simian
overlapping of canines, and he was able to grind his food by
moving his lower jaw from side to side in true hominid fashion.
The possession of this gap fails to make him an ape, but it also
renders his descent from the kinds of Australopithecines found in
South and East Africa unlikely. A second notable feature of P-4 is
that the roof of the palate is smooth, as in apes, instead of ridged,
as in men.
The Pithecanthropus Mandibles from the Djetis Bed
A piece of mandible found in the same level with P-4 is known
as Pithecanthropus B. It belonged to a different individual and is
just a little too small to match P-4’s palate. It consists of a frag-
2 G. Heithersay: “A Dental Survey of the Aborigines at Haast’s Bluff, Central
Australia,” MJA, May 30, 1959, pp. 721-9.
3 W. E. LeGros Clark: The Fossil Evidence for Human Evolution (Chicago:
University of Chicago Press; 1955), p. 94.
The Pithecanthropus Mandibles from the Djetis Bed 381
ment of the right branch, 82 mm. long, running from a break be-
tween the two right incisors almost to the gonial angle. It contains
all three molars and the sockets of the second incisor, canine, and
first premolar. It is a very large mandible, larger in nearly all di-
mensions than any other unquestionably human fossil jaw, and
Fig. 56 Mandibles of Meganthropus, Pithecanthropus B, and Wadjak 2.
A. Meganthropus (after von Koenigswald and Weidenreich ) ; B. Pithecanthropus
B (after Weidenreich); C. Wadjak 2 (after a cast). Meganthropus is represented by
fragniMits of two large mandibles found in Java in the Early Pleistocene Djetis
beds. This is the first specimen, found by von Koenigswald. It is larger than any
known human jaw and as large as those of Swartkrans and Kromdraai in South
Africa. It was probably that of an Asiatic Australopithecine, although some
anatomists consider it Hominine. The Pithecanthropus B jaw was its contemporary
Pithecanthropus B nearly matches the Pithecanthropus 4 skull. Between Pithe-
canthropus B and the next Australoid mandible, that of Wadjak 2, there is a gap of
over halt a million years. Wadjak 2 is a large, stout, human jaw with a chin but it
is as robust as that of Heidelberg. It also resembles that of Skhul 5 in Palestine—
the Australoid “Neanderthal” — very closely in form.
3 82 Pithecanthropus and the Australoids
certainly the stoutest. Its circumference holds the record to date.
Nevertheless, it is only two thirds as large as the Meganthropus
fragment found in the same deposit.
Although the actual symphysis (the sagittal line at the mid-
point between the two halves of the jaw) is missing, enough bone
is left in the center of the jaw to reveal the presence or absence of
a chin. It is absent. The angle of inclination (the angle between
the axis of the symphysis and the alveolar border) is about 58°,
the same as that of Meganthropus, and within five degrees of the
figures for the earliest Chinese and European mandibles, Sinan-
thropus and Heidelberg. Among living Australian aborigines this
angle runs to 750 and in many living Europeans it exceeds 90° —
a right angle.
Another fragment of jawbone from the same deposit is the so-
called Sangiran mandible, a piece 60 mm. long, again of the right
branch, and containing the first and second molars, a stub of the
second premolar, and the sockets of the first premolar and canine.
Going by the cast alone, Weidenreich described it, in 1945, as the
jaw of an orangutan. Meanwhile von Koenigswald had identified
it as a Meganthropus. In 1956 the latter wrote: “Not until later,
when I brought the original to New York and we were able to pre-
pare (clean) it better, could I demonstrate to him that the jaw
was human, while he convinced me that it was too different from
Meganthropus to belong to the same species. Since we had both
been mistaken, I named this new form of primitive man Pithe-
canthropus dubius.” 4
Most reviewers of the paleanthropic scene must also have con-
sidered this specimen doubtful, as they have not mentioned it. Yet
it exists. It is not, apparently, an ape, and it may be important.
(See Tables 38 and 39 for measurements of the bones and of its
two teeth.) The mandible is intermediate in size between Meg-
anthropus and Pithecanthropus B. Its angle of inclination is 540,
intermediate between the figure for the two jaws mentioned above
and those for living apes. How it fits into the hominid picture in
Late Lower Pleistocene Java remains to be seen.
4 Von Koenigswald: Meeting Prehistoric Man, p. 114. This name also, ap-
parently, contains a pnn on the name Dubois.
The Brain Case of the Infant Modjokertensis
383
The Brain Case of the Infant Modjokertensis 5
Also in the Djetis beds was found the faceless and baseless
skullcap of a baby a little under two years old. The bone is thin
(maximum = 3 mm.) but the tympanic plate, a part of the tem-
poral bone near the earhole on which the condyle of the lower
jaw moves, had begun to ossify prematurely, from the sapiens
Fig* 57 The Skulls of Homo erectus and Homo sapiens at the Age of Two
(Profiles drawn to the same scale). Above is the skullcap of the infant from
Modjokerto, Djetis beds, Java, associated with Pithecanthropus 4 and the Megan-
thropus jaw. Age, Late Lower Pleistocene. (After von Koenigswald. ) Below is the
skull of the infant from Starosel’e, Crimea, dated at Wiirm I ( 75,000-40,000 years
ago). The race is Caucasoid. (After Roginskil.) The age of the Modjokerto infant is
“a little under two years”; that of the Starosel’e baby, “eighteen or nineteen
months.’ The skull of the erectus baby is already elongated, its forehead is
sloping, the profile of the lambda region is straight, and the occipital bone
sharply curved. It already has brow ridges. The sapiens baby’s skull has a bulbous
forehead, a sharply bent parietal profile, and a more open occupital curve. This
illustration shows that erectus and sapiens skulls can be told apart as early as the
age of two.
5 Von Koenigswald: Meeting Prehistoric Man, pp. 72-82.
Clark: op. cit., p. 92.
384 Pithecanthropus and the Australoids
point of view. The forehead is more sloping than in modern babies
of the same age, the occipital bone more angular. Brow ridges had
just begun to sprout over the outer corners of the eye sockets, be-
hind which there was already a postorbital constriction.
This was, without doubt, a P-4 baby, in which the critical fea-
tures of Homo erectus had begun to show themselves at an early
stage of development. We are fortunate to have an erectus baby
of this age. When we come to diagnose the evolutionary status of
other baby skulls of later date, what we have learned from this
one will be useful.
Men of the Trinil Fauna 6
From the overlying beds of the Trinil fauna, also largely of
south Chinese origin, and from Trinil itself and its neighborhood,
three skullcaps have been taken — Pithecanthropus 1, 2, and 3 —
an inconsequential piece of mandible called Kedung Brubus, two
whole and two broken loose teeth, and, oddly enough, six femora.
P-i, the original Pithecanthropus erectus discovered by Dubois, is
a bare skullcap without face and with an incomplete base. Its esti-
mated capacity of 900 cc. matches that of Pithecanthropus 4, but
neither skull is complete enough for accurate measurement. P-2
is a smaller skullcap similar in all respects to P-i, and it has a
capacity of only 775 cc., a reasonably accurate figure because this
skull is the most nearly complete of the four.7 P-3 is a small piece
6E. Dubois: Pithecanthropus erectus, eine Menschendhnliche Vebergangsform
von Java ( Batavia, 1894 ) •
Dubois: “Figures of the Calvarium and Endocranial Cast, a Fragment of the
Mandible, and Three Teeth of Pithecanthropus erectus,” PKAW, Vol. 27, No. 5
(1924 b
Dubois: “Figures of the Femur of Pithecanthropus erectus,” PKAW, Vol. 29,
No. 9 (1926), pp. 1275-7.
Dubois: “The Sixth (Fifth New) Femur of Pithecanthropus erectus," PKAW,
Vol. 38, No. 8 (1935), PP- 850-2.
Von Koenigswald: “Neue Pithecanthropus Funde . . Meeting Prehistoric
Man.
Weidenreich: “The Skull of Sinanthropus pekinensis,” PSNSD, Vol. 10, ws 127
(1943).
7 The figure of 775 cc. is Weidenreich’s ( 1945). Von Koenigswald says 750 cc.,
and Boule and Vallois, 810 cc. (M. M. Boule and H. V. Vallois: Les Homines
Fossiles (Paris, Masson & Cie.; 1952), p. 120.
385
Men of the Trinil Fauna
of juvenile skull consisting of a fragment of occiput, and one com-
plete and one incomplete parietal bone.
These three skullcaps and pieces thereof are as thick as P-4, and
no larger. In P-i and P-2 the brow ridges are preserved. They are
not only massive, as befits Homo erectus, but also they have a
characteristic form. Seen in profile, they merge into the rest of the
frontal bone very gradually, with little depression. This may be
seen in later skulls from the same region and also among some liv-
ing Australian aborigines. It is quite different from the brow-ridge
profiles seen in other fossil lines, to be described presently. The
frontal bone is slightly keeled along its mid-line, and this keeling
continues along the sagittal suture, between the two parietal
bones. The keeling has nothing to do with a crest such as we have
seen in the larger Australopithecines. In fact, the temporal lines
are set relatively low down on the parietal bones.
Seen from above, the brow ridges form a nearly straight bar,
with no concavity in the profile line over the root of the nose, and
with relatively little backward curvature at either end, indicating
a relatively low figure for the frontal index of facial flatness. Be-
hind the brow ridges at either end is a deep hollow through which
heavy temporal muscles operated. This partial separation of the
brain case and facial skeleton is reminiscent of, but less extreme
than, the condition seen in Zinjanthropus.
The Trinil Pithecanthropus skulls (using Trinil to mean their
fauna ) differ from the earlier P-4 architecturally in that they are
shorter and narrower at the base. They are as broad as P-4 be-
tween the parietal bones, or a little broader, and equally high in
the vault or a little higher. These are changes of grade, not of line,
pointing in a modern, more fully human direction. During the
200,000 years or so which separate the two faunal levels, Djetis
and Trinil, evolution seems to have been at work, if not in increas-
ing brain size, at least in giving the container of this pilot organ a
less rugged and less bestial form.
The Kedung Brubus mandible is a small, triangular fragment
measuring about 35 mm. on each side, and including nothing in
the way of teeth except the root of a canine. As far as we can tell,
it could easily have been part of a Pithecanthropus jaw.
Incidentally, the Trinil faunal beds in some of the central Javan
386
Pithecanthropus and the Australoids
sites contain the earliest tools known in Java. They are crude but
clearly worked flakes. Because these flakes are technologically
well in advance of the crudest known choppers and chopping
tools, such as those found in Late Lower Pleistocene deposits in
Malaya, one must assume that Pithecanthropus had been a tool-
maker for some time before these flakes were made.
The Pithecanthropus Thighbones
I N 1891, a year after his discovery of P-i, Dubois also found a
complete human femur in the Trinil beds at the same site as the
skull. Some thirty years later he encountered five others, only one
of which was whole, in a box of old animal bones in his labora-
tory. The first femur has a curious bony growth, or exostosis, on
the inside of the shaft just below the neck. This femur — and the
others were like it — was long and slender, only moderately
curved, and had a pilaster ( a bracing ridge running down the
back of the shaft) of modern proportions. According to Loth,
however, the attachment for the adductor magnus muscle on the
underside of the shaft is a little higher than in modern femora.8
Except for this detail— in view of its length of 45.5 cm. — it could
have been the femur of a recently deceased Australian aborigine
or Papuan standing 5 feet 6 inches ( 168 cm. ) tall.
There is no reasonable doubt that these femora and the Trinil
skulls come from the same population because their fluorine con-
tent is identical, and it matches the fauna.9 Moreover, it is highly
unlikely that two kinds of men lived in Java during the Middle
Pleistocene, one race represented by four pieces from the neck up,
and the other by six pieces from the waist down. Pithecanthropus,
then, had the legs of an Australian aborigine and a skull that was
evolving in the same direction.
8 E. Loth: “Beitriige zur Kenntnis der Weichteilanatomie des Neanderthalers ”
Z FRK, Vol. 7 (1938), pp. 13-35.
9R. A. M. Bergman and P. Karsten: “The Fluorine Content of Pithecanthropus
and Other Specimens from the Trinil Fauna,” MKNA, Vol. 55, No. 2 (1952),
pp. 150-2.
The Teeth of Pithecanthropus
387
The Teeth of Pithecanthropus
Only twenty teeth that have been described may be at-
tributed to Pithecanthropus, four more than half the normal
dentition of an adult human being. Ten are in their sockets in
the maxillae of P-4, and two others were found loose in the same
site, and were probably his also as they fit two empty Sockets.
Three are imbedded in mandible B; two in the P. dubius mandi-
ble; and the other three, two molars and a premolar, were picked
up loose in the area of P-i. We will ignore the lower first molar
from the Sonde site, a probably modern tooth found on the sur-
face.
All but the three Trinil teeth are part of the Djetis Pithe-
canthropi. Of the three it is not certain whether the two molars
are human or belonged to an orangutan,1 and the premolar could
be modern.2 If these teeth were both human and contemporary
with the Trinil skulls, they indicate no dental evolution between
the Djetis and Trinil Pithecanthropus populations, and the lot
can be studied as a whole, with the above caveats borne in mind.
The only incisors attributed to Pithecanthropus are a right up-
per median and a right upper lateral found loose in the site of
P-4. They probably but not certainly had fallen out of the corre-
sponding sockets of P-4. Both seem to be within the modern hu-
man size range, as are the corresponding Australopithecine inci-
sors. Evidence of their shape is conflicting, and detailed descrip-
tions have not been published. The median incisor seems to have
one or more large tubercles at the base on the lingual surface, and
to have had raised edges. The lateral incisor appears to have had
raised edges. Moderate shoveling, such as these incisors seem to
. * W' K' Gregory and M. Heilman: “Further Notes on the Molars of Hespero-
prtheeus and of Pithecanthropus,” with an appendix by Gerrit S. Miller Tr. en-
titled Notes on the Casts of the Pithecanthropus Molars,” BAMN, Vol. 48,' Art.
13 (1923), pp. 509-30. Miller opted for the orang explanation;’ Gregory and
Heilman were uncertain.
2 Clark: op. cit., p. 51.
3 These descriptions are based on photographs that do not show details clearly,
e median incisor is depicted in von Koenigswald’s Meeting Prehistoric Man, on
page 59; the lateral in Weidenreich’s “Giant Early Man from Java and South
388
Pithecanthropus and the Australoids
have undergone, is found in 64 per cent of a composite series of
modern Australian aborigines,4 and the same condition has been
observed in Tasmanian teeth.
The two upper canines of P-4 are a little larger than the modern
maximum dimensions, and spatulate, as in man, rather than coni-
cal, as in apes. Apparently they overlapped the lower teeth a little,
but not enough to have prevented rotary chewing. The upper pre-
molars fall within the modern size range, but the upper second
premolar is unusually thick (in labiolingual width). The upper
first premolar also has three roots, like those of many apes and
most of the robustus group of Australopithecines, whereas the ear-
lier Australopithecines and most human beings have only two
roots for that tooth. A lower second premolar in the Pithecanthro-
pus B mandible again exceeds the modern range in both length
and breadth. A second specimen, the loose one from the Trinil site
that LeGros Clark considers of dubious age, has modern dimen-
sions.
Not counting those of P. dubius, we have six upper and three
lower molars. All of them, whether indubitably human or not,
have wrinkled enamel surfaces comparable to those of orangs and
of some Australopithecines. Their roots are long, stout, and di-
vergent. In P-4 the second upper molar is the largest; the first next
in size; and the third the smallest. In modern upper jaws the first
is usually the largest, followed in descending order of magnitude
by the second and third. Of the two upper molars from the Trinil
site, one is a second, the other a third. The third is larger than the
second, but this ranking is inconclusive because the teeth may
have come from two individuals, and may not even be human.
In general these teeth fall outside the modern range — including
that of the Australian aborigines — in labiolingual breadth more
than they do in mesiodistal length, and this is true of the denti-
tion as a whole. We are reminded, to a lesser degree, of the exces-
sive width of the Zinjanthropus teeth. However, two of the molars
China,” on plate 5-b. Von Koenigswald ( 1950, p. 59) called them both “extremely
shovelled, by far surpassing the condition seen in Sinanthropus,” a conclusion
that may refer to the development of tubercles but not to the morphology of the
dental borders.
4 A. Riesenfeld: “Shovel-shaped Incisors and a Few Other Dental Features
Among the Native People of the Pacific,” AJPA, Vol. 14, No. 3 ( 1956), pp. 505-22.
The Teeth of Pithecanthropus 389
are notably long and narrow; the lower third molar of Pithecan-
thropus B and the upper third of the Trinil trio.
As Weidenreich, following Selenka,5 pointed out, the degree of
wrinkling seen on the enamel of primate molars and the amount
of surface relief seen on the unworn cusps are inversely related.
Among the apes the orang has the most wrinkling and the lowest
cusps; the gorilla has the reverse. The heavily wrinkled third mo-
lars of Zinjanthropus are virtually cuspless. As one would expect
from this review, the Pithecanthropus molars have relatively low
cusps, which are not easy to identify from available publications.
The upper molars of P-4 have at least four cusps each, and the
upper second and third may have five. The three lower molars of
Pithecanthropus B all have at least five, and some may have a
sixth. The teeth are too worn for us to be certain. Of the ques-
tionable Trinil molars, the upper second is too worn to tell, and in
the upper third, which is highly wrinkled, the cusps are marginal
and hard to separate.6
As the first and second lower molars of the P. dubius mandibu-
lar fragment are close in size and form to those of Meganthropus,
Pithecanthropus, and the orang, these molars have been identi-
fied, by various authors, with all three.
On the whole, those teeth which we know definitely belonged
to Pithecanthropus because they were found in the sockets of un-
questionably human upper and lower jaws are much closer in size
to those of modern men than they are to those of the African Aus-
tralopithecines (except Telanthropus), Meganthropus, or Zinjan-
thropus. In a few respects they are more pongid than those just
mentioned, but as we have no lower first premolars, the compari-
son between the Pithecanthropus dentition and that of apes can-
not be carried to a conclusion. It is interesting that a man of Wei-
denreich’s stature as an anatomist should have been in doubt as to
whether the Sangiran molars belonged to a man or to an orang,
and that the team of W. K. Gregory and Milo Heilman, whose
combined competence in primate odontology was without peer,
should have had similar doubts about the Trinil molars. No one
has ever confused the Australopithecine teeth of Africa with those
s Weidenreich: “Giant Early Man . . . ,” pp. 64-8.
6 A. Hrdlicka : “Skeletal Remains of Early Man,” SMC, Vol. 83 (1930), p. 49.
390
Pithecanthropus and the Australoids
of chimpanzees or gorillas. The earliest hominids of Asia and its
fringing islands were more apelike dentally than their African
cousins. Moreover, the teeth of Pithecanthropus were closer to
those of Meganthropus than any of the African specimens of
Homo erectus yet found were to those of the most recent African
Australopithecines. Much more, however, remains to be discov-
ered in both the Oriental and the Ethiopian regions.
The Third Known Human Population of Java: Solo Man
I n t h e Solo River Valley of central Java may be seen, in various
places, a high terrace above the Trinil beds from which Pithecan-
thropus x, 2, and 3 were taken. This high terrace, in its Notopuro
beds, contains an abundant mammalian fauna belonging to a
phase of the Upper Pleistocene. Although most of the species are
still alive, the specimens recovered are larger than their modern
descendants. The horns of a buffalo, for instance, were extremely
long and widely spread. This and other evidence suggests that the
landscape was then more open and grassier than at present, for
now it is dense forest in which these prehistoric beasts woxdd have
difficulty moving about. Whether this indicates the somewhat
drier periods corresponding to the Third Interglacial, the major
Wiirm Interstadial, or some other time of reduced moisture is un-
known. From the standpoint of the skulls presently to be de-
scribed, an earlier rather than a later Upper Pleistocene date
would be more consistent with the evidence as a whole.
In this terrace, near a place called Ngandong, C. ter Haar, a
Dutch paleontologist, found over 25,000 pieces of mammal bone,
including eleven human calvarias and calvas, and two human
tibias. These finds were made between 1931 and 1933. The skulls
were all lying base upward and were in perfect condition. They
had not been moved or rolled. From each the facial skeleton had
been cut off, and in all but two the base had also been partially
removed, apparently by prying with a stick through the foramen
magnum, just as present-day Papuan head-hunters open up skulls
to eat the brains. In one, Solo 6 (S-6), the base was whole, and in
another, S-11, only the orbital roof had been broken out. One may
Sex, Age, and Injuries of the Eleven Skulls 391
surmise that these skulls had been carried to the river bank for a
feast of brains. But, then, why was S-6 left unshelled, why were
all the faces cut off elsewhere, why were they all lying neatly up-
side down, and why had the tibias been brought along? These leg
bones had not been split for marrow.
Neither the dating nor the face- and brain-removing technique
can be explained by the accompanying artifacts. Twenty- two
stone tools, as yet undescribed, were found.' The discoverer
picked up, with the skulls, a sting-ray barb, far from its home in
the sea. Such barbs are used by modern Australian aborigines. He
also found a notched bone implement, presumably a spear point,
and a piece of antler that may or may not have been used.
Sex, Age, and Injuries of the Eleven Skulls
Table 37 gives the vital statistics of the eleven skulls, except for
a small fragment of what may be a twelfth specimen adhering to
S-3. Only six were whole enough for detailed measuring, as shown
on Table 17. These six, S-i, 5, 6, 9, 10, and 11, are all adult. Two
are called male, three female, and in one case Weidenreich, who
studied them, lefused to commit himself. His determination of
sex, which he himself questioned at various times, was based
largely on size and on the thickness of the bones of the cranial
base.
In skulls S-i, 3, 5, 9, and 10, all the sutures are completely fused.
This condition is rare in modern man, however aged, and few
primitive men live to ripe old ages. One infers an earlier fusion of
sutures in Solo than in modern man. Furthermore, the skulls are
all two or three times as thick as modern crania, and just as thick
as their Middle Pleistocene predecessors.
That a thick skull had a survival value in Soloese days is sug-
gested by the roster of bone injuries in this series. Wholly aside
from the post-mortem mutilations, scars left from nonfatal bat-
tles are prevalent. On S-4 a large lesion shows where a heavy, and
‘ Von Koenigswald: “Der Solo-Mensch von Java: ein Tropischer Neanderthaler,”
in von Koenigswald, ed.: Hundert Jahre Neanderthaler (Utrecht: Kemink en Zoon;
1958), pp. 21-6.
392
Pithecanthropus and the Australoids
TABLE 17
THE NGANDONG SKULLS
Sex
Age
Parts Present
S-l
probably f.
mature adult
Calvarium, base missing from nasion to
mastoids, warped
S-2
?
child, 3-5 yrs.
A frontal bone
S-3
m.
mature adult
Calva: both parietals and adjacent parts of
frontal and occipital
S-4
probably f.
adolescent
Calotte, frontal and both parietals
S-5
m.
mature adult
Calvarium, complete but for broken base
S-6
f.
young adult
Calvarium, complete, base whole
S-7
f.
adolescent
Fragment of right parietal
S-8
m.
adolescent
Two detached matching parietals
S-9
?
mature adult
Calvarium, most of base missing
S-10
f.
mature adult
Calvarium, most of base missing
S-ll
m.
young adult
Calvarium, front of base missing
probably also sharp, implement pierced the scalp and outer
bone table. The lesion then healed. S-6 bears a similar scar. S-i
and S-io each carried to death a square injury in which the diploe
had been laid bare. S-i also showed additional minor scars. All
four heavily battle-scarred victims are called females and at least
two of them must have been injured while young, because they
died young. Their social life seems to have been active, and S-i
may have been particularly popular.
The Racial Anatomy of the Ngandong Skullcaps
From Mynheer ter Haar, who found them, the skulls passed to
W. F. F. Oppenoorth,8 who published articles about the six first
discovered, then to Ralph von Koenigswald, who delivered all but
S-g to Franz Weidenreich in New York. S-9 had been presented to
8 W. F. F. Oppenoorth: “The Place of Homo soloensis Among Fossil Men,”
in G. G. MacCurdy, ed.: Early Man (Philadelphia: J. B. Lippincott Company;
1937), PP- 349-6o.
The Racial Anatomy of the Ngandong Skullcaps 393
Emperor Hirohito during the Japanese occupation of Java and Lt.
(now Dr.) Walter A. Fairservis, Jr., liberated it from an imperial
collection in Kyoto and returned it to the company of its fellows.
At the time of his death in 1948, Weidenreich was still writing his
definitive report of them.9
This document was published just as he left it, ending in the
middle of a sentence. It does not give the over-all dimensions of
each skull, but he had earlier published a series of measurements
taken on casts,1 and more recently Ronald Singer has published
some of the measurements of the originals 2 (see Table 37). These
figures, combined with a family likeness evident in the photo-
graphs, indicate that the Solo skulls belonged to a single popula-
tion.
The lowest cranial capacity, 1,035 cc'> exceeds the highest for
Pithecanthropus, 900 cc. by a considerable gap, which might be
closed had we enough specimens of each to represent a popula-
tion statistically. The highest, 1,255 cc-> well within the central
three fourths of the modern human range. If Weidenreich’s de-
termination of sex is right, the three females have an average ca-
pacity of 1,042 cc., and the two males of 1,158 cc.; and on this
basis the undesignated S-9 ought to be classed as male. The cra-
nial capacities of the two Trinil Pithecanthropi, P-i, male, and
P-2, female, differ by 125 cc., and those of modern male and fe-
male Australians by over 150 cc. These comparisons make the sex
difference of 110 cc. postulated for Solo man reasonable.
Thus, we can suggest a male sequence as follows: Pithecantho-
pus = 900 cc.; Solo = 1,150 cc.; living Australians = 1,350 cc. The
female sequence would be: P = 775 cc.; S = 1,040 cc.; Austra-
lians = 1,180 cc. This exercise in numbers places Solo almost ex-
actly in the middle of the procession in both sexes, a grade ( or a
half grade, if one prefers ) above Pithecanthropus and below the.
aborigines.
9 Weidenreich: “Morphology of Solo Man,” introduction by Von Koenigswald.
APAM, Vol. 43, Part 3 (1951). See also Weidenreich: “Giant Early Man from
China and Java,” APAM, Vol. 40, Part 1 (1945).
1 Weidenreich: “The Skull of Sinanthropus pekinensis,” pp. 111-16.
2 Ronald Singer, in von Koenigswald: “Der Solo-Mensch von Java: ein Tropi-
scher Neanderthaler,” in Hundert Jahre N eanderthaler (Utrecht: Kemink en
Zoon N. V.; 1958), p. 22.
394 Pithecanthropus and the Australoids
The cranial lengths of this series exceed both those of Pithe-
canthropus and those of the modern Australian aborigines, al-
though the internal brain length is actually intermediate. Kap-
pers, who invented an index between cranial length and the
length of the endocranial cast ( brain length ) , gives the following
figures: P— 1 = 83.7; S— 5 = 83.97; Sinanthropus = 83.6; and
among the living Australians, males = 88.5 and females = 90. 19. 3
This places Solo man in the same grade as the Trinil Pithecan-
thropi as far as the development of bony crests is concerned.
The lateral crests were equally robust. Weidenreich gives two
cranial breadths, the usual biparietal or outer brain-case width,
and a so-called bicristal, which is taken lower down, across the
mastoid crests. In modern men, the biparietal is almost always the
greater of the two, but in Pithecanthropus and Solo it is the
smaller. In Solo the lower dimension (the bicristal) remains
comparable to that of Pithecanthropus, but the brain-case width
increases a good ten millimeters, and both basion-bregma and
auricular head heights increase about fifteen millimeters. Thus,
the increase of 150 cc. in brain size from Pithecanthropus to Solo
involved a parallel growth in each of the three basic cranial di-
mensions. The size of the brain case changed more than its
shape did.
Seen from the side, the Solo profile is still Pithecanthropoid.
The brow ridges still slope gradually back from glabella without
the abrupt nick found in some other fossil men, notably Sinan-
thropus in China and Steinheim in Europe. This feature has been
noted in some modern Australian crania, as well as in living
aborigines. Again like Pithecanthropus, the Solo skulls have a dis-
tinct, projecting nuchal crest which forms the posterior landmark
of the entire skull. The area of temporal muscle attachment is bor-
dered not by faint lines but by raised crests. These borders follow
the same contour as the skull itself, and although the enclosed
area of temporal muscle attachment is adequate for a powerful
jaw, they do not swing far upward as in the Australopithecines
and to a lesser extent in Sinanthropus. Solo shares this feature
with Pithecanthropus.
3 C. U. Ariens Kappers: “The Endocranial Casts of the Ehringsdorf and Homo
Soloensis Skulls,” JAiuit., Vol. 71, Part 1 (1936), pp. 61-76.
The Racial Anatomy of the Ngandong Skullcaps 395
In four of the skulls, S-i, 5, 6, and 11, either a stub of the nasal
bones or a scar on the frontal, indicating the place where the two
join, is left. These traces show that the nasal bones came down
almost straight from the frontal without the pronounced nasal
notch present in many living Australian aborigines and Euro-
peans.
Seen from front or rear, the skull looks angular and ill-filled, its
profile broken into planes. This is both a Pithecanthropoid and an
Australoid trait. In most living races, as in most fossil men, the
profile is rounded. The base of the skull is nearly flat, both inside
and out, but the back part of the occipital floor slopes gently up-
ward from the level of the middle of the foramen magnum to the
nuchal crest. Indeed, the foramen magnum itself has two levels,
•TABLE 18
DIMENSIONS OF THE
HYPOPHYSEAL FOSSA*
S-11
Modern Men (after Pruitt)
L =
22 mm.
10.7 mm. (range = 6-16)
B =
22 mm.
10.0 mm. (7-17.5)
Depth =
9 mm.
8.7 mm. (3-15)
* After Weidenreich, 1951.
horizontal to the eye-ear plane in front and inclined slightly up-
ward behind. As in Pithecanthropus, the mastoid processes are
large.
Inside the foramen magnnm of S-11, Weidenreich found that
the hypophyseal fossa, or sella turcica, which is the seat of the
pituitary gland, was exceptionally large, with between three and
four times the volume of the same cavity in modern crania. This
discovery implies that Solo man had a very large pituitary and
that therefore the endocrine balance of this population was dif-
ferent from that of Homo sapiens. From the behavioral viewpoint
this is a very important observation. Washburn and Howell, how-
ever, have challenged Weidenreich ’s interpretation of the anat-
omy of S-n’s cranial base, and the matter cannot be settled with-
out further study of the skull, which is in Bandung, Indonesia.4
4 S. L. Washburn and F. C. Howell: “On the Identification of the Hypophyseal
Fossa of Solo Man,” AJPA, Vol. 10, No. 1 (1952), pp. 13-22. The authors claim
396
Pithecanthropus and the Australoids
Fig. 58 Profiles: Australoids from Trinil to Niah. A. Pithecanthropus
2 from Trinil; B. Solo 11; C. Wadjak 1; D. the youth from Niah Cave,
Borneo. Pithecanthropus 2 is shown here because number 1 is not available in
profile view and numbers 3 and 4 are fragmentary. It is the smallest of the four,
with a cranial capacity of only 775 cc., only slightly over the Australopithecine
range. Its profile is characteristic of the Pithecanthropi as a group, with angular
lines and brow ridges which join the forehead with little depression. The Solo
skull is essentially the same but larger. The tops of the nasal bones of Solo 11 are
preserved below the brow ridges. They come directly from the frontal, with little
notching. Wadjak 1, the earliest sapiens skull from Java (with Wadjak 2), closely
resembles that of an Australian aborigine. The skull from the Niah Cave in North
Borneo, that of a youth of sixteen, establishes the existence of Australoid Homo
sapiens in that island 40,000 years ago. (Drawings A and B after Weidenreich,
1951; C after Weidenreich, 1945, and a cast; D after Brothwell and Higgs, 1961.)
The Face of Solo Man
Whoever removed the faces of the Solo skulls did a thorough
job. All that is left is the upper borders. The brow ridges, although
large, do not form a solid bar, as in the Trinil skulls, but are di-
vided. Seen from above, they form a Cupid’s bow, less projecting
that what Weidenreich called the hypophyseal fossa was really the sphenoid sinus,
broken open from above. Yet in 1947 Weidenreich described the internal anatomy
of S-11 in detail. Weidenreich: “Some Particulars of the Skull and Brain of Early
Hominids . . . ,” AJPA, Vol. 5, No. 4 (1947), pp. 387-428.
The Ngandong Leg Bones 397
at the center than over each eye. From the front they also appear
to constitute a double arch. The outer ends, which are particu-
larly thick, bend slightly downward. The center also dips down-
ward and the nasal bones take off from the frontal at a low level,
without invading that massive bone at all. The lines marking the
sutures between the frontal, lachrymal, and nasal bones lie nearly
on the same level. Here there is no angularity; the region of the
mask was broad and rather flat. Tentative measurements of the
upper breadth of the nasal bones and of the interorbital distance
in four skulls, S-i, 5, 6, and 11, show that the nose was broad at
the root, and the eyes set far apart. On Solo 11 it is possible to
measure the internal biorbital chord and its subtense to nasion,
and to calculate, with these measurements, the frontal index of
facial flatness. The figure is 15.0, matched today only by Mongo-
loids and approximated by Negritos and Bushmen. This particu-
lar Solo man had a very flat face, at least in the region of the
eyes and the root of the nose.
When we compare the Solo skulls with the Djetis and Trinil
Pithecanthropi, we find that the three sets of bones complement
each other, and that they show a well-marked continuity over a
period of as much as 400,000 years, in a nearly constant envi-
ronment, and at a slow rate of evolution for man. This environ-
ment was the wet tropics, in its time an outer corner of the
earth.
The Ngandong Leg Bones
Like Pithecanthropus before him, Solo man left parts of his legs
for posterity, but they were different parts from those of the Trinil
Pithecanthropi. Instead of six femurs we have two tibias. Tibia A
is a piece of shaft 30 cm. long broken off at both ends. It was once
at least 12 cm. longer. Tibia B, which is nearly complete, is 36.5
cm. long. These tibias are straight, unflattened, and, like the
Pithecanthropus femurs, modern. If A was a male, as seems
likely, he could have been 5 feet 10 inches tall ( 178 cm. ) or a lit-
tle shorter if his shins were disproportionately long. If B was a
398 Pithecanthropus and the Australoids
woman, she could have been 5 feet 4 inches tall (162 cm.).5
These statures fall within the range of living Australoids and far
exceed that of the living Javanese.
What Name, Mr. Solo?
I N Oppenoorth’s initial description of the first six Ngandong
skulls, he proposed the taxonomic title Homo ( Javanthropus )
soloensis. This was in 1932. Later in the same year he dropped the
parenthetical Javanthropus, leaving only Homo soloensis. In 1937
he said: “In my first publication I proposed to unite H. soloensis,
H. rhodesiensis, H. wadjakensis, all proto-australian forms, into a
separate subgenus, Javanthropus, but — and I completely agree
with Dubois that they all belong in this group — that name was not
well chosen and it is better to drop it. Yet we have in Homo
soloensis the oldest at present known representative of Homo
sapiens fossilis.” 6
Meanwhile, in 1934, von Koenigswald had dubbed the skulls
Homo neanderthalensis soloensis. When Weidenreich wrote his
final monograph he said : “Earlier studies led me to the conviction
that Ngandong man is not a true Neanderthal type but distinctly
more primitive and very close to Pithecanthropus and Sinan-
thropus. For this reason I ranked Solo man with the same group
of early hominids as the two later forms and called the whole
group Archanthropines. . . . Considered from this point of view,
it is entirely irrelevant whether Solo man is called Javanthropus
soloensis or Homo soloensis. I decided to use simply ‘Solo man.’ ” 7
From the post- 1960 point of view, this historical discussion
seems as irrelevant to me as it did to Weidenreich in the late
1940’s. Solo man was closely related to the two successive Pithe-
canthropus populations. He had nothing to do with Neanderthal;
and as we shall see in the next chapter, he occupied the same
5 The least diameters of tibia A are 28 mm. anteroposteriorly and 30 mm.
bilaterally; it is also 46 mm. thick at the top where broken. The comparable fig-
ures for tibia B are: 25 mm., 25 mm., and 40 mm.
6Oppenoorth: “The Place of Homo soloensis . . . pp. 358-9.
7 Weidenreich: “Morphology of Solo Man,” p. 227.
The Fourth Known Human Population of Java 399
evolutionary grade as Sinanthropus. Although larger-brained
than his predecessors, he was still Homo erectus.
The Solo-like Brain Case from Aitape, New Guinea
In all past and present Australoid regions, no Homo erectus
skulls have yet been found outside of Java, with one possible ex-
ception. That is a brain case unearthed in 1925 at Aitape, in the
Finsch Coast area of northeast New Guinea.I * * * * * * 8 Its date is Pleisto-
cene, and as the dating was carried out by oil geologists working
with fossil molluscs, there is little possibility of error. What part of
the Pleistocene it came from we do not know, but, on other evi-
dence, we may suppose that it was Upper Pleistocene and late,
dating from a time when New Guinea and Australia were con-
nected by the Sahul Shelf, and Wallacea contained more dry land
than it does today.
The specimen is a calva of a female aged about forty-five years,
and consists of most of a frontal bone and portions of both parie-
tals. It has very heavy brow ridges, a sloping forehead, and an
angular cranial contour. The temporal lines follow the profile of
the brain case. This specimen is not only clearly Australoid, but it
bears a striking resemblance to the Solo calvaria. Whether it had
crossed the erectus-sapiens line we cannot say without further
study.
The Fourth Known Human Population of Java:
Wadjak Man
I n 1890, before he discovered the famous Trinil skullcap, P-i,
Eugene Dubois found two other fossil skulls at a place called
Wadjak in central Java. They had been cemented in breccia, and
were unaccompanied by implements. Because the fauna was mod-
ern in every sense, they were probably later than the Solo skulls
and late Pleistocene in date, at the earliest. As the site has since
been destroyed by quarrymen, the exact date may never be
known. For thirty years Dubois kept the world in ignorance of
8 F. J. Fenner: “Fossil Skull Fragments of Probably Pleistocene Age from
Aitape, New Guinea, RSAM, Vol. 6 ( 1944), pp. 335— 54.
400 Pithecanthropus and the Australoids
these to him relatively uninteresting skulls, and failed to describe
them until 1931.9 Luckily, we have casts of both specimens.
Wadjak 1, or W-i, is a nearly complete cranium, the base of
which is largely intact, and a piece of mandible. In the cranium
several considerable breaks and gaps may be seen in the right
temporal and occipital regions, and both zygomatic arches are
gone. Only seven upper teeth are in situ and unbroken: all five
premolars and molars on the right side, and the second and third
molars. An eighth tooth, which could be a canine, is stuck in its
matrix in the roof of the palate. At the time of casting (my de-
scription is based largely on a cast ) , the skull had not been fully
cleaned.
The piece of mandible is also still breccia-bound. It consists of
85 mm., more or less, of the right branch, running from the socket
of the first molar to the gonial angle, the rear half of the first mo-
lar tooth, and the second and third molars. The ascending ramus
is broken off just above tooth level. Most of the lingual surface and
the bottom edge remain to be cleaned.
W-2, while more fragmentary, is in better shape, and has no ad-
hering matrix. It consists of five pieces, which could be assembled
as adequately as Weidenreich assembled those of Pithecanthro-
pus 4. They are : ( 1 ) a piece of frontal including the brow ridges,
with the upper part of the nasal bones, and with part of the left
zygomatic bone extending under and defining the left eye socket;
(2) the maxillae, including the palate and the floor of the nasal
passages, extending 25 mm. up the side of the nasal opening ( this
is a little more than the corresponding piece of P-4) and contain-
ing, in situ, all the permanent upper teeth except the incisors and
the right first molar; (3) most of the occipital bone; (4) part of
the right temporal, including the mastoid and the ear hole; and
(5) the mandible, the right half of which is nearly complete. The
left half is broken off 10 mm. behind the level of the third molar.
All the permanent lower teeth are present except the right ca-
nine, both first premolars, and the right third molar.
9 Dubois: “The Proto-Australian Fossil Man of Wadjak, Java,” PKAW, Vol. 23,
No. 7 (1921), pp. 1013-51.
G. Pinkley: “The Significance of Wadjak Man . . . ,” PNHB., Vol. 10, No. 3
( 1936), pp- 183-200.
The Wadjak Brain Cases
401
Dubois, who considered Wadjak 1 a female on grounds which
could be contested, gave it a cranial capacity of 1,550 cc. As the
interior of the skull had not been cleaned, this figure was appar-
ently obtained by a formula based on modem skulls. In the light
of more recent knowledge, we may use the von Bonin formula for
Australoid skulls. The result is a figure of 1,475 cc-> which may
still be too high in view of the massiveness of the bones. Wadjak 2,
which Dubois classified as a male — probably correctly — may have
had a brain of more or less the same size.
Despite these corrections, the Wadjak pair are large skulls; in
size they fit within the upper fourth of the modern Australian
range for males, and as far as brain size goes, they are fully
evolved members of the species H. sapiens. This places them one
brain-size grade above the Solo skulls, which in turn are a grade
above Pithecanthropus. With this exercise, we have passed
through more than 400,000 years of time, from the most primitive
known human form to a man with a modern brain size, all within
a small section of a single, medium-sized island having a mini-
mum of environmental change and probably little if any advance
in technology.
For the first time in the Pithecanthropus-Australoid sequence
we have a population represented by the essential parts of two
faces. Moreover, this is the first time since the Djetis Pithecan-
thropi that mandibles are available. The face, teeth, and jaws of
the Wadjak specimens can be expected to reveal some progressive
changes from the features of their predecessors, but not as many
as brain size will indicate. It is unlikely that feeding habits, which
involve the jaws and teeth, kept pace during this journey
through time with advances in human relations, which are more
closely linked to the growth of the brain.
The Wadjak Brain Cases
O f t h e two Wadjak brain cases, only W-i can be described as
a whole. It is a large skull in all three principal dimensions, com-
paring favorably with any in the world. It differs from Solo prin-
cipally in height and in that the bicristal and biparietal breadths
I
402 Pithecanthropus and the Australoids
are apparently the same; this is difficult to determine because the
crests are broken and the brain case is both broken and warped.
The length is not from crest to crest but from a moderately promi-
nent glabella to a rounded occipital bulge. The two heights, ba-
sion-bregma and auricular, differ by over 20 mm., whereas in the
earlier Javanese skulls the differences are only from 12 to 15 mm.
This reflects a change in the morphology of the brain case.
In 1947, Weidenreich observed that the truly primitive brain
cases, including those of Pithecanthropus, Sinanthropus, and
Solo, differed from those of modern men in two related respects.1
In the more primitive types, the upper profile of the brain is
nearly flat; in the more advanced ones it is humped. In the more
primitive skulls, the floor of the brain case is virtually flat, while
in the more advanced ones it is bent downward at both ends, as
well as in the median sagittal line. This is caused by the growth of
the parietal lobes of the brain in the center, and by the growth of
the middle lobe of the cerebellum, which is concerned with vol-
untary movements of the muscles, particularly those of the limbs,
including the fine movements of the hands in skilled work. Among
other effects, this bending lowers the base of the brain relative to
the position of the ear hole. It is hard to observe this phenomenon
in W-i because of breakage, warping, and poor cleaning, but in
the occipital bone of W-2 the concavities that hold the base of the
cerebellum and the occipital lobes of the cortex are cupped in-
ward and downward. Wadjak man’s brain was not only curved
on top but it also had acquired the bending specified in Weiden-
reich’s study. In these respects as well as in absolute size his brain
was more evolved than that of Solo.
Morphologically, the Wadjak brain cases resemble those of
Pithecanthropus and Solo in a familial way. There are still planes,
although the contours are more nearly round, the mastoids are
still large, and the temporal muscle lines still follow the contour
of the skull roof. The forehead is sloping and extends upward and
backward from glabella without a marked depression. Although
the zygomatic arches are gone, enough is left of the zygomatic
1 Weidenreich: “Some particulars of skull and brain of early hominids and
their bearing on the problem of the relationship between man and anthropoids,”
AJPA, Vol. 5, No. 4 (1947), pp. 387-428.
The Wadjak Mandibles 403
process of the left malar of W-i to indicate a bizygomatic diame-
ter above the modern Australian maximum.
The Wadjak Faces
W-i has a large face. It is long, broad, and flat. The brow ridges
are heavy only over the nasal region, and the nasal bones take off
from the frontal without much depression, but with more than we
have seen previously. The nose is exceptionally flat, the interorbi-
tal distance is wide, and the orbits wide and low. Although W-i’s
face is too battered, bruised, and poorly cleaned to permit accu-
rate measurements of the chords and subtenses needed for the
first, third, and fourth indices of facial flatness, I have ventured to
try the second, or simotic index, which expresses the degree of
flatness of the saddle of the nose. It is 24.7 (?),2 which, if correct,
places W-i in the company of Negritos, Negroes, Bushmen, and
Hottentots, and gives her or him a slightly flatter nose than the
means for living Papuans and Melanesians. Because Wadjak 2’s
upper facial region is in better condition, it is possible to calculate
the first index, the frontal index of facial flatness. This is 18.6, right
in the middle of the range of means for recent Australian aborig-
ines and Tasmanians and inside that of Papuans and Melanesians.
In W-i the alveolar protrusion is tremendous, and its appear-
ance is exaggerated by the flatness of the upper part of the facial
skeleton. In both specimens the nasal aperture is very broad and
its border guttered, as in Pithecanthropus 4. The palates are
long, broad, and deep, within the metrical ranges of modern Aus-
tralian crania in all three measurements, but definitely smaller in
every way than our only previous specimen from this area, Pithe-
canthropus 4.
The Wadjak Mandibles
Our first virtually complete mandible in the Pithecanthropus-
Australoid sequence is that of Wadjak 2. What is missing on one
side is present on the other. Because the palate of W-2 does not
2 The question mark indicates that the figure is not certain, because of break-
age.
404 Pithecanthropus and the Australoids
fit it perfectly, it is apparent that the palate has been widened in
the region of the right second and third upper molars in back of a
break line. This distortion has been allowed for in the figure for
palate breadth in Table 38.
The first thing that one notices about this mandible is that it
looks modern because it has a chin, but after a little handling and
matching with other mandibles a second fact becomes clear — this
is a very large, heavy jawbone. It is just as large and heavily built
as Pithecanthropus B. In other words, in the Pithecanthropus-Aus-
traloid line as seen in Java, the lower jaws remained equally large
and strong for over a half million years. The Wadjak 2 mandi-
ble is also just as large and heavy as the famous Heidelberg jaw
from Germany, which is at least 360,000 years old. But morpho-
logically the Wadjak 2 mandible is more advanced than either
Pithecanthropus B or Heidelberg in that it has a chin.
The Wadjak Dentition
In the upper jaw there are two, three, or four specimens of
every tooth except the incisors. In general, these upper teeth are
as long as those of P-4 anteroposteriorly, but narrower labiolin-
gually. The indices of robusticity ( length times breadth ) are thus
lower. In particular, the second upper molars of both Wadjak
specimens are smaller in every way than those of P-4. Although
the Wadjak upper teeth run a little larger than the modern Aus-
tralian mean, they are well within the modern range. Also, for the
first time we find the modern size sequence in upper molars, in
which the first molar is the largest and the third the smallest.
In the lower jaws there is at least one tooth for each place in the
row. What has been said above about the sizes of the upper teeth
in comparison to other specimens and series is equally true of the
lowers. In the Wadjak 1 mandible the molar sequence is also the
same as in the upper jaw, but in the Wadjak 2 mandible the
third molar is larger than the second.
Among the Australian aborigines the cusp number for each
molar tooth is usually ^ ^ that is, all three upper molars have
5 5 5
The Significance of Wadjak
405
four cusps, and all three lowers have five. In Wadjak 1 it is -4~ ~3,
5-4-5
and in Wadjak 2 it is -5_4_4.3 Both Wadjak x and Wadjak 2, then,
5 4 5
had fewer cusps on their molars than any of the Pithecanthropus
specimens; and Wadjak l’s molars were a little more reduced,
that is, advanced, than those of the average modern Australian
aborigine, whereas those of Wadjak 2 are identical with aboriginal
teeth.
Only the teeth of Wadjak 2 are well enough reproduced in pho-
tographs or casts to permit the study of other details. Each upper
canine has a teatlike ridge running up the center of the lingual
side, reaching from the base of the crown to the level where the
crown is worn (about 6 mm.) Comparable but narrower ridges
are found on the lingual sides of the upper left canine and on all
four lower incisors. On the upper left canine, the edges of the
tooth are raised. These features are common in Australian teeth.
The Significance of Wadjak
The study of the two Wadjak specimens completes our series
of four consecutive populations that lived in the center of one of
the smaller islands of central Indonesia during the Pleistocene.
This is but a tiny fraction of the total area presumably occupied
by members of the Pithecanthropus-Australoid line during a pe-
riod of over a half million years. It shows continuity, variability,
and evolution. Homo erectus was still alive there less than a hun-
dred thousand years ago, and Homo sapiens appeared there ten
thousand years ago or earlier. Sometime between Solo and Wa-
djak, the transition was made.
But it was not necessarily made in Java itself. The Djetis Pithe-
canthropi, the Trinil Pithecanthropi, Solo, and Wadjak may rep-
resent successive invasions from a center of Australoid evolution
somewhere in the north, such as Siam or Indochina, where there
have been no Duboises or von Koenigswalds to seek out fossil
men. If Java was a periphery of southeast Asia, Australia is a pe-
3 Pinkley: The Significance of Wadjak Man”; and also observation of casts.
406 Pithecanthropus and the Australoids
riphery of Java. Bearing in mind the principle that the outer pe-
ripheries of zoogeographic regions tend to be inhabited by archaic
kinds of animals, let us see what Australia has to offer in the fos-
sil-man line.
Fossil Man in Australia
We do not know when human beings first began to bother the
kangaroos by appearing in Australia, but it was undoubtedly later
than man’s first appearance in any other continent, except Ant-
arctica. The first people to reach North America had a broad
Pleistocene highway to walk over. The level of the earth’s oceans
controlled their time of passage. The first to reach Australia and
New Guinea had no such dry road, for the islands of Wallacea
rise steeply from the sea. With or without the presence of the
Sunda and Sahul shelves, whoever made the crossing still had to
hop from island to island on rafts or small boats, and the greatest
distance that had to be traversed was about 50 miles.
When the great shelves were above water, migrants could enter
Australia by way of Timor as easily as they could get to New
Guinea via the Moluccas. When the shelves were submerged, New
Guinea was the only feasible port of entry. Which route was used
first we do not know, nor are we sure that the Timor-Sahul Shelf
route was used at all, but the present distribution of racial traits
in the entire Australian faunal area favors the latter.
The only concrete evidence favoring a late Pleistocene date of
entry is the age determination of the Aitape find, which is still an
isolated discovery and needs confirmation. The oldest Carbon-14
date yet obtained from an archaeological site is 6,740 ± 120 b.c.,
for Cape Martin, South Australia,4 associated with an archaic cul-
ture known as the Tartangan. Because a still more primitive cul-
ture, the Kartan, has been found in several sites below the Tartan-
gan, Tindale estimates that the Kartan must have begun at least as
early as 9,000 b.c.5 No evidence is yet available which indicates
4 D. J. Mulvaney: “Australian Radio-carbon Dates,” Antiquity, Vol. 35, No.
137 (1961), pp. 37-9-
5 N. B. Tindale: “Ecology of Primitive Aboriginal Man in Australia,” in
A. Keast, R. L. Crocker, and C. S. Christian: “Biogeography and Ecology in
Australia,” MB, Vol. 8 (1959), pp. 36-51.
The Keilor Skull
407
that entry took place before 10,000 b.c., the very time when Mon-
goloid peoples had, as we shall soon see, begun pressing into
southeast Asia out of China.
Linguistic evidence suggests that the dispersal of the Austra-
loid peoples occurred less than 20,000 years ago: as explained in
Chapter 1, languages lexically related to one another can be no
older. All Australian languages are mutually related. The Papuan
languages probably belong to the same family; so apparently does
Andamanese (the languages of the Andaman Island Negritos)
and even, it has been claimed, the Mon-Khmer languages of
southeast Asia, which are spoken by Mongoloids, and of parts of
India, where they are spoken by both Mongoloids and Austra-
loids.
Whatever the archaeological and linguistic evidence may
prove, we have a number of mineralized human remains from
Australia, and more effort has beeen spent in discussing their ages
than in describing them. Of the lot only three have been tenta-
tively accepted by the profession as having any antiquity: Keilor,
Talgai, and Cohuna.
The Keilor Skull
I n 1940 a fossil skull was found in a sandpit in a place called
Keilor, ten miles north of Melbourne. It is heavily mineralized and
a Carbon- 14 date for the same terrace, but not the same site, is
6,55° ± 25° B-c- (W-169), well within the range of the Cape
Martin date. The importance of this skull lies not in its age but in
its close resemblance to Wadjak 1, which Weidenreich found to
be within the range of twins.6 This resemblance has been some-
what reduced by new measurements of the Wadjak- 1 cast, but it
is still there. If not identical twins, they could have been brothers.
Keilor was an adult, and apparently a male, but this is not cer-
tain.
G Weidenreich : “The Keilor Skull: a Wadjak Type from Southeast Australia,”
AJPA, Vol. 3, No. 1 (1945), pp. 21-32.
J. Wunderley: “The Keilor Skull, Anatomical Description,” MNMM, No. 13
(1943), pp- 57-70.
W. Adam: “The Keilor Fossil Skull, Palate and Upper Dental Arch,” MNMM,
No. 13 (1943), pp. 71-8.
408 Pithecanthropus and the Australoids
The published cranial capacity, 1,593 cc-> is greater than the top
of the Australian range. When recalculated with the formula used
for Wadjak 1, it reduced to 1,464 cm., which is a more realistic
figure, close to the new figure for Wadjak 1. Keilor’s face was of
moderate length, his nose flat, his orbits low. He had the same
Negrito-Bushman look seen in Wadjak 1. This settles the question
whether people with this kind of face could have been in any way
ancestral to the more pointed-faced, living aborigines.
The Talgai Skull 7
The Talgai skull was found in 1884 at Darling Downs,
South Queensland. It lay seven feet down in undisturbed clay un-
der black soil, near the top of the clay. The skull eventually
reached the University of Sydney, where it was described in
1918, but one upper left median incisor tooth remained in Queens-
land because its owner refused to sell it. The skull was heavily fos-
silized and badly crushed. Shortly after death, however, someone
had removed the front part of the base in the usual way and pre-
sumably for the usual purpose. It was apparently a male, four-
teen to sixteen years old. The skull is smaller than Keilor, with a
cranial capacity of about 1,300 cc., and it is as low-vaulted as
Solo. Although its brow ridges are moderate, it has a nuchal crest,
and altogether it is a more primitive specimen than Keilor. The
palate is as large as that of Wadjak 2, and nearly square; the mo-
lar-premolar rows are nearly parallel, and the canines and inci-
sors form nearly a straight line across the front. In this respect it
resembles the older Pithecanthropus 4, although there is no evi-
dence of a diastema.
As neither wisdom tooth had erupted, Talgai had died young,
but he had lived long enough to wear the crowns of his first molars
down to the dentine. His teeth are larger than those of either of
the Wadjaks or of Keilor, and approach those of Pithecanthropi 4
and B in size. In fact, his canine is a shade wider mesiodistally
than P-4’s, but it is not as thick labiolingually. The premolars and
7 S. A. Smith: “The Fossil Human Skull Found at Talgai, Queensland,” TRSL
(B), Vol. 208 ( 1918), pp. 351-87.
The Cohuna Skull
409
first molar are nearly as large as P-4’s but his second molar is con-
siderably smaller, as in modern skulls.
Despite his big teeth, Talgai did not resemble the Javanese
skulls facially. Metrically, the skull falls at about the middle of the
modern Australian range in details of the nasal skeleton and or-
bits. Morphologically, it is not very flat-faced, but has a depressed
nasion and a rather beaky profile. Its features are run-of-the-mill
Australian. If it and Keilor were contemporary, the facial fea-
tures of the Australian aborigines were variable at that time, as
they are today.
The Cohuna Skull 8
The third and last on our list of moderately well authenti-
cated Australian skulls of some antiquity is Cohuna, found in
1925, two feet below the surface of the soil, in the Murray River
Valley on the edge of Kow Swamp, Cohuna, Victoria. The skull
was completely mineralized and filled with silt. Nearby were
about fifty pieces of equally fossilized skulls and bones which
have not been described, and eleven recent aboriginal skeletons.
This skull was mutilated twice. Immediately after death some
fellow-aborigine broke off the base to get at the brain. Thousands
of years later a modern, white Australian who owned the skull cut
from it two perfect circles of bone, each 27 mm. in diameter. As
one of these circles included inion, it is difficult to reconstruct
the nuchal border of the occipital bone. Although the cranial ca-
pacity has not been published, the von Bonin formula for Austra-
loid skulls places it well over 1,500 cc. This figure is probably
much too high because the forehead is exceedingly low and slop-
ing, and the auricular head height of i22(?) mm. represents a
peak rather than a plane. Without doubt, the true capacity of this
skull falls within the modern Australian range.
Aside from the extreme slope of the forehead, which rivals that
of the Pithecanthropi except that it continues up farther, and its
8 N. W. G. Macintosh: “The Cohuna Cranium, History and Commentary from
Nov. 1925 to Nov. 1951,” Mankind, Vol. 4, No. 8 (1952), pp. 307-29.
D. J. Mahony, W. Baragwanath, F. Wood-Jones, and A. S. Kenyon: “Fossil
Man in the State of Victoria, Australia/* RIGC , (16th) Washington, 1936, pp.
1335-42-
410 Pithecanthropus and the Australoids
very heavy brow ridges, this skull is notable for its tremendous
prognathism, and its wide bizygomatic diameter, which exceeds
the modern Australian range. Furthermore, the palate was not
horseshoe-shaped but rectangular. Unfortunately we have no
measurements of the teeth. Although its face is extremely long, its
features, like Talgai’s, are Australian. Its nasal root is deeply sunk
under the brow ridges, and its facial profile is not flat.
The Pithecanthropus- Australoid Line
These three skulls, Keilor, Talgai, and Cohuna, whatever their
ages, definitely link the living Australian aborigines to the succes-
sion of four Pleistocene populations in Java, and to the Aitape
brain case from New Guinea. Dubois first saw this connection and
Weidenreich agreed with him. It has been more recently accepted
by Boule and Vallois, and by Piveteau. Pithecanthropus of course
is extinct, and so is Solo man, but neither died without progeny.
Their extinction took the well-known form of evolution by suc-
cession, or, more simply, of evolving into something else.
When the ancestors of the Australian aborigines and their is-
land-dwelling neighbors arrived at their present homes, they were
already marginal people; and after they had settled there they
fell into a marginal geographical pattern of their own. Among the
living aborigines, for example, the curliest hair and the heaviest
brow ridges are found on the peripheries of the continent and its
offshore islands. Whether this implies a succession of invasions, a
slow trickle of genes from the northwest during the period of
Mongoloid pressure on southeast Asia and Indonesia, or local
evolution radiating from a central Australian focus, we do not
know, nor is the question pertinent to the thesis of this book.
Some evolution has probably been taking place, but, as one
would expect in a marginal area of the Southern Hemisphere,
its over-all rate cannot have been rapid. One still finds recent abo-
riginal female skulls with cranial capacities of 930 cc., 946 cc., and
956 cc. whose owners apparently met the demands of their cul-
ture well enough to live to maturity; 9 and I have myself measured
9 Macintosh: op. cit.
4n
The Mapa Skullcap
a living married woman named Topsy (see Plate XXXII) with a
head length of 184 mm., a breadth of 121 mm., and an auricular
height of iog mm., flesh included. Her cranial capacity was under
1,000 cc. and she was a normal member of Tiwi society, described
in Chapter 3.
This study leads to several conclusions. One is that the Austra-
lian aborigines are still in the act of sloughing off some of the ge-
netic traits which distinguish Homo erectus from Homo sapiens.
Another is that, as rates of evolution differ in different parts of
the world, populations belonging to a given evolutionary grade in
different places cannot be closely related if their life spans are
hundreds of thousands of years apart. Having established a base
line of evolutionary tempo for the Pithecanthropus-Australoid
line, we can, in subsequent chapters, see how it matches the rates
of other lines.
Human Evolution North of Java in the Pleistocene
Hav 1 n g left two areas of light, Java and Australia, we now
enter a realm of virtual darkness. The skeletal history of the more
northerly parts of the Eastern Oriental region is extremely frag-
mentary, consisting almost entirely of a few scraps of bone, a
handful of teeth, preliminary notices of two or three newly discov-
ered finds, and that is all. Only two finds can be definitely called
Pleistocene, those of Mapa in south China and of the new skull
from the Niah Cave of Borneo.
The Mapa Skullcap
The older of the two is the Mapa calva, probably that of an
adult male, found in the province of Kwangtung, south China.1
Its date is late Middle or early Upper Pleistocene, later than ei-
ther the Djetis or the Trinil Pithecanthropi and probably earlier
than Solo. It consists of the frontal, parietal, and nasal bones and
the lower border of the right eye socket. As it bears certain resem-
1 Ju-Kang Woo: “Fossil Human Skull of Early Paleanthropic Stage Found at
Mapa, Shaoquan, Kwangtung Province,” VP, Vol. 3, No. 4 (1959), pp. 176-82.
412
Pithecanthropus and the Australoids
blances to both Solo and Sinanthropus, it can best be studied after
we have described the latter in the following chapter. It is men-
tioned here merely to set the northern boundary of the area we are
talking about, at the time it lived.
The Upper Pleistocene Skull from Niah Cave,
North Borneo
I N *959 Tom Harrison, who had been excavating the vast and
famous cave at Niah, North Borneo, for several years, discovered a
human skull at a depth of eight feet four inches. It was associated
with chopping tools and large, coarse flakes, comparable to the
Soanian of India, and it has been given a Carbon- 14 date of 39,-
600 ± 1,000 years ago by the Groningen Laboratory.2 This makes
it possibly as old as Solo and certainly older than Wadjak, and
about as old as the first Upper Paleolithic men of Europe.
It is the skull of a youth between fifteen and seventeen years
old, probably female, definitely sapiens, and equally definitely
Australoid. Its closest resemblance is to the skulls of modern Tas-
manians. It has very little brow-ridge development, a deep nasal
root, and a vertical forehead. The parietal bones have high, promi-
nent bosses; the occiput is well rounded; and the mastoids are
small. The palate and teeth are smaller than those of the fossil
Australian skulls but comparable in size and shape to those of liv-
ing Australians and Tasmanians. The face is short, the nose broad.
The tracks of the meningial arteries on the inside of the parietal
bones are modern in complexity and form.
This skull indicates that, by the time of the Gottweig Inter-
stadial in Europe, or about 38,000 b.c., the Australoid subspecies,
at least in Borneo, had crossed the threshold between Homo erec-
tus and Homo sapiens, either through local evolution alone, or
as a result of gene flow from the Mongoloid region to the north.
2W. G. Solheim II: “The Present Status of the ‘Paleolithic’ in Borneo,” AP,
Vol. 2, No. 2 of 1958 (Hong Kong, i960), pp. 83—90. The C-14 date number is
GRO-1338.
D. R. Brothwell: “Upper Pleistocene Human Shull from the Niah Cave, Sara-
wak,” S MJ, Vol. 9, No. 15-16 ( i960), pp. 323-49.
The Mesolithic-N eolithic Transition in Indonesia
413
The Mesolithic-Neolithic Transition in Indonesia
Following the time of the Niah skull, we must jump at least
25,000 years until the end of the Pleistocene, and later. The main
seat of the Australoids then shifted from Indonesia and southeast
Asia to the Australian region, whereas in Indonesia and southeast
Asia Australoids were gradually replaced, except in a few margi-
nal refuges, by Mongoloids. We would like to know the details of
this replacement. When, for example, did the ancestors of the
Negritos shrink to their present size? Did they do this in one or
several acts of shrinking? When did the Mongoloids come in? Did
they first arrive as food gatherers, and in later waves penetrate as
agriculturalists, or was the first wave already agricultural?
We cannot answer these questions satisfactorily because the ex-
isting skeletal material is scanty, and what little has been found
has been inadequately described. It seemed less important to its
discoverers than the older, fossilized skulls and bones, and it is
mostly fragile and hard to handle. Also the archaeological se-
quences in this area have not yet been fully clarified and co-ordi-
nated. The term Neolithic does not in itself distinguish between
food-gathering and food-producing cultures but seems to include
some of both. Finally, it is hard to tell which of the different
Mesolithic” and “Neolithic” tool industries evolved locally and
which were derived from outside.
About the same time that he discovered the Wadjak skulls,
Dubois also found a skeleton in a cave near the ancient lake of
Wadjak. It was less fossilized than the Wadjak bones, and cov-
ered with red ochre. Dubois identified it as Mongoloid.3 It was
round-headed. In another cave at Bodjonegoro, also in Java, in
what may have been a Mesolithic deposit, although it could be of
later date, P. van Stein Callenfels 4 found some scraps and bits of
human skeletal material. Of these, three molar teeth were meas-
ured. They are too big for Negritos, and a little too big for Java-
3 Dubois: “The Proto- Australian Fossil Man of Wadjak, lava,” PKAW, Vol. 2
No. 7 ( 1921 ) .
4 Van Heekeren: The Stone Age of Indonesia (The Hague: Martin Nijhoff;
1957), PP- 78-9.
4]4 Pithecanthropus and the Australoids
nese, who for small people have large teeth. They fit easily into the
Pithecanthropus-Wadjak size range.5
In the nearby cave of Sampoeng, in a series of “Neolithic” de-
posits, Callenfels and his associates found a number of burials.
W. A. Mijsberg was able to restore one cranium, called Sampoeng
F ,B He measured this cranium and its teeth carefully. In every one
of ten cranial dimensions, and in eight of nine cranial indices, it
falls within the modern Javanese range as determined from ca-
daver material in Batavia. The one exception is the upper facial
index, in which it exceeds the range, but this index depends to a
certain extent on diet, because heavy chewing spreads the zygo-
matic arches and increases the face breadth. As the teeth also
match those of the Batavian cadavers in size, we may well believe
that this skull belonged to an early Mongoloid ancestor of the
Javanese, whether agricultural or not we do not know.
To the east, in Celebes, two energetic Swiss cousins, Paul and
Fritz Sarasin, discovered in 1902 a tribe of backward folk living
partly in caves. They took various measurements of these people,
who were called Toala, and excavated the floor of the caves they
lived in, discovering some artifacts and a few human bones.7 As
a result of this research, they became convinced that the Toala
were close kin of the Veddas, a hunting people still living in Cey-
lon, who are actually small and primitive Caucasoids. They drew
this conclusion both from their study of the living and from their
examination of the skeletal remains exhumed from the floor of
the cave. This identification snowballed during the last half cen-
tury, until the presence of Veddoids in Indonesia, and in Malaya
as well, became textbook dogma. So it has remained, although
0 Left upper first molar is 12.3 mm. in anteroposterior length and 13.1 mm. in
labiolingual breadth. The right upper first molar is 12.8 mm. by 13.4 mm.; a lower
third is 13.0 mm. by 12.5 mm.
6 W. A. Mijsberg: “Recherches sur les Restes Humains de Goewa-Lawa a
Sampoeng et des Sites Prehistoriques a Bodjonegoro (Java),” in Hommage du
Service Archeologique des lndes Neerlandaises au Premier Congres des Prehis-
toriens d’ Extreme-Orient a Hanoi, 26-31 Jan. 1932 (Ratavia: Societe Royale des
Arts et Sciences; 1932).
7 Paul and Fritz Sarasin: Reisen in Celebes (Wiesbaden: C. W. Kreidel; 1905);
Versuch einer Anthropologie der Insel Celebes (Wiesbaden: C. W. Kreidel;
1905)-
The Mesolithic-Neolithic Transition in Indonesia 415
Mijsberg refuted this identification before 1950.8 He measured the
living and found them no different from the other more or less
Mongoloid inhabitants of Celebes, particularly similar to their
neighbors the Buginese. He measured also some sub-Recent
skeletal material from the cave of Bolabatu, Lamontjong Cave
(excavated by the Sarasins), Panganrejang Cave, and Lompoa
Cave.
From Bolabatu came one calvarium and a mandible. Available
measurements for both are within the Buginese range, and the
minimum frontal forehead breadth of 98 mm. is too great for a
Vedda, and so is the mandibular height of 31 mm. One female
skeleton from Lompoa Cave was four feet eight inches ( 142 cm. )
long, which is in the Pygmy range, but many living Indonesian
women are equally short. The teeth are also small, smaller than
those of living Buginese, but within the Javanese range.
This material, on the whole, indicates that the people who lived
in the Toalian caves in geologically Recent times were similar to
the living inhabitants of the region. The deposits in the cave in-
clude Bronze Age artifacts, and also “Neolithic” implements that
are called Toalian. Whether the people whose skeletons Mijsberg
measured were food gatherers or slash-and-burn cultivators is not
known.
Moving on to Sumatra, we come upon a report that “a few
fragments of skulls, hardly sufficient for a final racial determina-
tion, were found in one of the shell heaps of north Sumatra. . . .
Watsl . . . came to the conclusion that they showed Papuo-
Melanoid racial characters.” 9 Finding Papuo-Melanesian charac-
ters in sub-Recent but prehistoric bones was a favorite sport in
southeast Asia at that time. We cannot be sure that the Suma-
tran cranial scraps did not belong to Negritos.
In the Philippines, a Negrito skull was found, before 1921,
8 D. A. Hooijer: “Man and Other Mammals from Toalian Sites in S. W. Cele-
bes,” PKAW, Sec. 2, Vol. 46, No. 2 (1950), pp. 1-164, especially 59-74.
9R. Heine-Geldern: “Prehistoric Research in the Netherlands Indies,” in
P. Honig and F. Verdoorn, eds.: Science and. Scientists in the Netherlands Indies
(New York: Chronica Botanica Co.; 1945), pp. 129-67. After J. Watsl: “Prahis-
torische Menschenreste aus dem Muschelhiigel von Bindjai-Tamiang in Nord Su-
matra,” in a Festschrift: Otto Reche zum 60. Geburtstag . . . ( Munich-Berlin,
1939), pp- 237-43-
41 6 Pithecanthropus and the Australoids
under ten feet of alluvial deposit below the Rio Pasig in Manila.1
This is the first Negrito skull we have unearthed, but unfortunately
it cannot be dated.
Mesolithic and Neolithic Remains from Indochina
With a characteristic interest in archaeology, the French exca-
vated widely in Indochina during the last quarter of the nine-
teenth century and well into the 1930’s, stopping only with the
advent of World War II. They found three principal post-
Pleistocene cultural levels: a Mesolithic, then an early Neolithic
characterized by stone axes polished on the cutting edge only,
and finally a late Neolithic with axes polished all over. In all three
periods, the fauna was the same as that of today, except that no
bones of domestic animals have been found. Exactly when agri-
culture came in we do not know, but it is difficult to understand
why these people needed the vast numbers of axes they made un-
less they were clearing garden patches, or at least ringing the
bark of trees in preparation for burning.
Up to 1938, a total of thirty-five skulls intact enough for study
had been found and more or less described.2 Almost all come from
Laos, although some are from Tonkin. This region is close to the
Chinese border. Most of the remains were found in grassy country
well over 3,000 feet high, on the main invasion route of Mongoloid
peoples into Indochina. Not only was this highroad extensively
used in historic times; it is even serving this purpose today. The
Miao tribes of southern China advanced a full six degrees of lati-
1 D. Sanchez y Sanchez: “Un craneo humano prehistorico de Manila (Fili-
pinas),” MRSE, Vol. 11 (1921).
P. Huard and E. Saurin: “£tat Actuel de la Craniologie Indochinoise,” BSGI,
Vol. 25, No. 1 ( 1938), pp. 1-104.
2 Huard and Saurin: op. cit.
J. Fromaget and E. Saurin: “Note Preliminaire sur leS Formations Cenozoiques
et Plus Recentes de la Chaine Annamitique Septentrionale et du Haut Laos,”
BSGI, Vol. 22, No. 3 ( 1936), pp. 1-48.
H. Mansuy: “Contribution a 1’fitude de la Prehistoire de l’lndochine: V. Nou-
velles Decouvertes dans les Cavemes du Massif Calcaire de Bac-Son (Tonkin);
VI. Stations Prehistoriques de Keo-Phay, de Lai-Ta, et de Bang-Mac, dans le
Massif Calcaire de Bac-Son (Tonkin); VII. Neolithique Inferieur (Bacsonien) et
Neolithique Superieure dans le Haut-Tonkin,” BSGI, Vol. 12, Nos. 1, 2, 3 (1925).
Mesolithic and Neolithic Remains from Indochina 417
tude southward over it in the century that ended in the 1930’s.
If we are to find the earliest Mongoloid skulls in southeast Asia,
this is the place to look for them.
Luckily, an early Mongoloid skull has been found there, at Tam
Pong. It belonged to a young adult female about twenty years
old, who had not yet cut her wisdom teeth. A stature of about
five feet two inches (157 cm.) has been calculated from her
long bones. The skull is of modern size, with a capacity of over
1>35° cc- It is delicate in structure, well rounded, has no brow
ridges but does have well-developed mastoids. The orbits are not
as high as in most Mongoloids, but the interorbital distance is
great, and the nasal bones lie flat. Although the nasal opening is
wide, the bones themselves are very narrow at the top, and con-
stricted below nasion. The face is long and wide, and the malars
(cheekbones) are salient below the lower orbital margin. There
is no canine fossa. The palate is large and parabolic, the chin
well developed. There is no prognathism.
The skull is certainly neither Australoid nor Negrito. It must
then be either Caucasoid or Mongoloid. The shape of the orbits is
Caucasoid, but the structure and protrusion of the malars, the
length of the upper face, the shape and flatness of the nasal bones,
as well as the guttering at the border of the nasal opening, are all
Mongoloid, in a general sense. I am satisfied with the conclusion
that Mongoloid food gatherers had begun to enter southeast
Asia from the north fairly early in postglacial times. As Laos is a
northern frontier country, this evidence does not indicate how far
south these Mongoloids penetrated at what times. That they did
not immediately replace the earlier peoples can be seen from a
study of the Lower, or Early, Neolithic skeletons.
From the site of Tam Hang come four adult skeletons of the
Lower, or Early Neolithic, the time of partially polished axes and
no domestic animals. Two are males and two females. Both fe-
males were pregnant. The males are numbered S-3 and S-5, the
females S-2 and S-4.
Starting with the female S-2, we note that although most of
the face is missing, the mandible is present. This is a remarkably
short skull, but the breadth and height dimensions make up for
3 Fromaget and Saurin: op. cit.
4i 8 Pithecanthropus and the Australoids
its deficiency in length, giving it a cranial capacity of about
1,230 cc. The skull is thin and infantile, with a straight forehead;
the mandible modern and tiny. Without reasonable doubt, this
woman was a Negrito, resembling the modern Andamanese in
cranial structure. Her stature, four feet eleven inches, or 150 cm.,
lies on the upper border of the Negrito range.
Her companion, the male S-3, was an inch shorter, or 147 cm.
tall; and his cranial capacity, 1,430 cc., was greater. This skull is
practically complete. It is delicate, well rounded, somewhat bulb-
ous in the forehead, with an open metopic (frontal) suture, small
malars and a feeble development of the zygomatic arches, a
sharp-bordered nasal opening, a straight chin, and no progna-
thism. He, too, was a Negrito.
S-5, the other male, has no cranial base, and most of the face
is gone, but the mandible is there. The cranial capacity is about
1,340 cc. The skull is narrower than S-2 or S-3, and its orbits
seem to be higher, although it is difficult to tell because they are
incomplete. Unfortunately, we do not know how tall it was. From-
aget and Saurin suspect that it is not fully Negrito, and they may
be right.
S-4, our second female, is nearly complete. The skull is broader
and higher than the others from the same site, and it has a much
more massive face. Its malars project forward, but not as much as
in the Mesolithic skull from Tam Pong, which it resembles in
many ways, including the fact that it has high orbits and alveolar
prognathism. Her stature of five feet one inch, or 155 cm., is too
great for an ordinary Negrito woman. This individual may repre-
sent a Mongoloid-Negrito mixture, if not a full Mongoloid.
In sum, this little group of Early Neolithic people interred in
the site of Tam Hang were Negritos — who had begun mixing with
Mongoloid peoples from the north — Mongoloids of the same
general type as the Mesolithic woman of Tam Pong. The Negritos
were characterized by small stature; relatively long lower legs
and lower arms; a delicate, somewhat infantile skull form that is
rounder and higher than anything we have seen before; and small
faces.
In the same site, at the base of the Upper or Late Neolithic, a
skullcap was found in a mutilated state. It had apparently been
Mesolithic and Neolithic Remains from Indochina 419
used as a cup or bowl and is a very long, narrow calva, with a
cranial index of 70, a figure we have not encountered since deal-
ing with fully Australoid specimens. It thus represents a third
element in the Indochinese population; we shall presently discuss
this element more fully.
Higher up in the Late Neolithic stratum in the Tam Hang site
were found a group of one adult and two juvenile skulls. Only one,
that of an eleven-year-old, is complete enough to study. With a
cranial index of 78, it is mesocranial; its forehead is a little bulb-
ous, its nasal root smooth, its face rather small even for its age,
but it has alveolar prognathism and shovel incisors. In general it
seems to combine the elements of the Mesolithic and Early
Neolithic specimens of the neighborhood.
Other sites contain the skeletons of local groups quite different
from those we have just seen. We find in them a new element, al-
ready anticipated by the presence of the drinking-bowl calva at
Tam Hang. Nine skulls from Lang Cuom and one from Dong
Thuoc belong to this type, which the French call Melanesian,
although they qualify a few as Indo-Melanesian and Australo-
Melanesian. This new element is, simply, a very long and nar-
row-headed skull, with an index in the low seventies, which is
also high-vaulted, with the height equaling or exceeding the
breadth.4
The skull looks like an Australian aboriginal skull with some of
the corners rounded off and the brow ridges and other bony
struts toned down. The face also resembles a softened, less primi-
tive-looking, Australian face. The relationship to the Australian
type is clear. The reduction in primitive features could be due
either to mixture with something else or to evolution.
Personally I prefer the evolutionary explanation, at least as the
major cause of this change, although mixture with both Negritos
and early Mongoloids cannot be ruled out. Still, neither of these
two would give these modified Australoids their extreme cranial
form. Moreover, it seems to me a little premature to call them
Melanesian, because that implies a Negroid hair form. We know
4 Some of the skulls from Lang Cuom and elsewhere were badly deformed by
earth pressure, which reduced their breadths greatly. Cranial indices in the low
sixties in this series should be discounted.
420 Pithecanthropus and the Australoids
nothing of the hair form of any prehistoric skull. The Melanesians
of today are believed to be Papuans modified by fairly recent
Polynesian admixture. If the skulls of Lang Cuom and elsewhere
resemble those of Melanesians, this simply means that a mixture
which took place in the Pacific islands also occured in Indochina
as an independent phenomenon. In south-central India there live
several million primitive agricultural people who speak languages
of an Indochinese type. These tribesmen, the Munda, Ho, and
Santal, are a mixed group, with three phenotypes predominating
to a certain extent in individuals. Most numerous is an evolved
Australoid, next a Mongoloid, and least a Negrito or Negroid. If
the ancestors of these peoples came from Indochina, it is difficult
to believe that the majority had curly hair.
Six of the skulls at Lang Cuom were more or less Mongoloid,
as were two from Pho Binh Gia and one from Keo Phay. Two of
the Lang Cuom skulls were also credited with mixed Negrito
features. Whatever the accuracy of these diagnoses, it is clear that
the interesting mixture which we still see in south-central India
was already in process of formation. Another conclusion is that
the transformation of the countries of southeast Asia from an Aus-
traloid realm to a southern extension of the Mongoloid lebensraum
had not been completed by the end of the Lower or Early Neo-
lithic; and the cranial material from the Upper or Late Neolithic
is too scanty to indicate whether the great rush took place then or
later. One suspects that most of the replacement occurred in the
full light of history, after Alexander the Great had met his end,
when the Chinese empire had begun its great expansion, which
has not yet ended, and after the Chinese had started to squeeze
“barbarian” Mongoloid peoples like the Thai and Shan and Laos
out of their cool mountains onto the steaming plains of the south.
Migrating in tightly organized, Iron Age tribes, they pushed the
aborigines before them.
Bypassing Siam, which has to date produced not a single pre-
historic human bone, because no one has really looked, we make
our next stop at a kitchen-midden near Guak Kepah, Wellesley
Province, Straits Settlements, where van Stein Callenfels found a
mandible in 1935. The date is Mesolithic or sub-Recent. Mijs-
Prehistoric Populations of the Western Oriental Region 421
berg, who studied it, considered it similar to the jaws of modem
New Caledonians.5 He was apparently impressed by the resem-
blance between certain stone discs with double perforations found
at the site and similar ornaments used until recently in New Cale-
donia. As the New Caledonian mandibles are Australoid, this
one may be called Australoid too, in a general sense.
Six skulls unearthed by I. Evans in various Malayan caves
and rock shelters, notably Gunong Sennyum, Lenggong, Gunong
Pondok, and Bukit Chuping, were studied by Duckworth at Ox-
ford.6 They are called “Neolithic,” mostly late, because stone tools
were used well into the Metal Age, and lack faces. All of them
follow a single pattern, that of the “Melanesians” of Lang Cuom,
except that at least one, from Sennyum, is more rugged, with old-
fashioned heavy brow ridges, and looks more like the Australoid
prototype. We can skip Burma, which is as lacking in prehistoric
crania as Siam, and push on to India.
Prehistoric Populations of the Western Oriental Region
Today the eastern half of the Oriental faunal region is in-
habited almost entirely by Mongoloids. Yet the evidence that we
have just reviewed indicates that the ancestors of the Burmese,
Thais, Indochinese, Malays, and Indonesians arrived in their pres-
ent homes quite late. The movement southward out of China be-
gan in the postglacial Mesolithic, reached its peak in historic
times, and is still taking place in the sense that the Chinese them-
selves are moving into southeast Asia.
The earlier inhabitants of southeast Asia and Indonesia are rep-
resented today by a few enclaves of Negritos and Australoids and
by the food-gathering Mongoloid tribes, such as the People of the
Yellow Leaves in Siam, the Kubu in Sumatra, and the Punans in
5 Mijsberg: “On a Neolithic Palae-melanesian lower jaw found in a Kitchen-
midden at Guak Kepah, Province Wellesley, Straits Settlements,” PTCPFA, 1938
(Pub. 1940), pp. 100-8.
6 W. L. H. Duckworth: “Human Remains from Rock-Shelters and Caves in
Perak, Pahang, and Perlis and from Selinsing,” JMBR, Vol. 12, Pt. 2 (1934),
after Huard and Saurin.
422 Pithecanthropus and the Australoids
Borneo. There is no evidence to indicate that these culturally
primitive Mongoloids arrived in their present homes before the
postglacial Mesolithic, and it is possible that some of them are
feral, that is, refugees from agriculture. There are no primitive
Caucasoids in the area, and no evidence exists that there ever
were any.
When we cross the Burmese mountains into India, which geo-
graphically includes Pakistan and Ceylon, we encounter a com-
parable but different situation. Here the majority of the popula-
tion, including speakers of both Indo-European and Dravidian
tongues, is Caucasoid. A minority of the Dravidian speakers, com-
posed mostly of tribal peoples outside the caste system, are Aus-
traloid, and a very few are Negroid, but both have probably taken
over Dravidian speech from their culturally more advanced
neighbors.
India also shelters some Mongoloids, as for example the Khasis
and Garos of Assam, but none of them are food gatherers and
there is no reason to suppose them to have entered India before
the Neolithic. The marginal, casteless groups of food gatherers,
comparable to the Negritos, Sakai, Andamanese, and so on farther
east (the Andaman Islands are politically a part of India) be-
long to three races: Negrito or Negroid, Australoid, and Cauca-
soid. The Negrito or Negroid element is always found in mixture
with Australoid, whereas the Caucasoid tribes are usually un-
mixed. In central India there also live some agricultural tribes
that speak three languages, Ho, Miinda, and Santal, related to the
Mon-Khmer speech of Burma and Indochina. These people are
Australoid with Mongoloid admixture. At least the Mongoloid
element probably came in from the east in the Neolithic or later.
In southeast Asia and Indonesia the earliest population was
probably Australoid in the wide sense; the Negritos and Aus-
traloids who survive there today can only be descended from this
ancient polymorphic, and probably regionally variable, popula-
tion pool. As India is also a part of the Oriental region, it seems
logical to suppose that the same is true there also, but we cannot
test this hypothesis adequately because we have little skeletal ma-
terial from the subcontinent, and none of it is ancient.
The oldest we have is seven skeletons from a site called
Prehistoric Populations of the Western Oriental Region 423
Langhnaj near Gujarat in West Pakistan, in the Indus Valley.7
Buried in a habitation site with mesolithic implements and with-
out pottery, they are probably older than the Bronze Age civiliza-
tion of that region. Four have been partly described; three are
called males and one a female. They were moderately tall people.
One male was five feet seven inches in height ( 170 cm.), and one
female five feet four inches ( 162 cm. ) . Their lower arms and shins
were moderately long compared to their upper arms and thighs,
and they were of slender build. Their skulls are long and narrow,
and the men had sloping foreheads, but at least one woman had a
bulbous one. Owing to earth pressure, their facial bones are dis-
torted. Nevertheless, one woman seems to have been prognathous,
particulai ly in the upper jaw, and one man had a Caucasoid-
looking lower nasal skeleton. These people were either Caucasoid
or Australoid, or most likely a combination of both.
As the skeletal material from the Bronze Age civilization of the
Indus Valley includes Caucasoid, Australoid, and Mongoloid
skulls, all we know is that these three subspecies were represented
in northwest India as early as 2400 b.c.8
The circumstantial evidence of geographical distribution
slightly favors the greater antiquity of the Caucasoids, because
of the racial situation in Ceylon. That island was settled bv
the ancestors of the Singhalese, who came from northern India,
speaking an Indo-European tongue, about 500 b.c. Later, Tamil-
speaking people from south India settled the northern part of the
island. Both these peoples are primarily Caucasoid, although the
Singhalese also contain a Mongoloid element.
When the Singhalese arrived, they found the island occupied
by two groups of primitive hunters, which they called Yakkhas
and Nagas.9 Sometime between the arrival of the Singhalese and
the period of European exploration and colonization, either one
of these groups wiped out the other or they fused into a people
called Veddas. The living Veddas are Caucasoid. However, they
' I. Karve and G. M. Kurulkar: “Human Remains Discovered So Far,” in H. D.
Sankalia and Karve: Preliminary Report on the Third Gujarat Prehistoric Expedi-
tion (Bombay: Times of India Press; 1945).
8 The skulls are in Calcutta, the postcranial bones in Karachi. I have examined
the skulls, which have not been completely described in publication.
N. D. Wijeskera : The People of Ceylon (Colombo: Gunasena; 1949), p. 32.
424 Pithecanthropus and the Australoids
are divided into clans, some of which are said to have light and
others dark skins. Perhaps uniquely among simple food gatherers,
they recognize some clans as noble and others as servile. These
distinctions tend to substantiate the observations of the early
Singhalese, that the aborigines consisted of more than one people.
Recently Paul Deraniyagala, the director of the museums of
Ceylon, and a well-known paleontologist, found six skeletons in
southern Ceylon associated with a Mesolithic stone industry.1
These skeletons have been given a Carbon- 14 date of 110 b.c. ±
200 years, which places them nearly four hundred years after the
arrival of the Singhalese. No trade goods were found with the
burials as might be expected were this date correct. As the graves
were shallow, the charcoal samples may have been contaminated;
a somewhat earlier date would make more sense.
One male skeleton had a stature of five feet six inches ( 167 cm. ) ;
a female skeleton was five feet one inch ( 154 cm. ) tall. In a series
of 138 living male Veddas measured in the 1930’s by J. R. de la
H. Marrett, the mean stature was five feet one and a half
inches, or 156.78 cm., and the range was from 134 to 172 cm.2
These two Ralangodans, although taller than most modern Ved-
das, were within their stature range.
Although from the published measurements it is not possible to
calculate cranial capacity, this capacity is probably close to that
of the Veddas, whose mean is 1,260 cc. for 138 males. The teeth
also are of moderate size, comparable to those of living Singhalese,
who are Caucasoids. I have been unable to find a published ac-
count of Vedda teeth, but I have seen them on skulls and in the
living, and am sure that they are at most only as large as those of
the Singhalese. On present evidence I cannot state that the
Ralangodan teeth differ from those of the Veddas.
In 1957 I saw the Balangodan skulls then in Ceylon and in
i960 I inspected the one labelled T-24-B which Dr. Deraniyagala
had left at the American Museum of Natural History in New York.
1 P. E. P. Deraniyagala: “The Races of the Stone Age and Ferrolithic of Cey-
lon,” JRAS (Ceylon Branch), Vol. 5, Pt. 1 (1956), pp. 1-23.
Deraniyagala: “An Open Air Habitation Site of Homo sapiens Balangodensis,”
SZC, Vol. 28, Pt. 2 (1958), pp. 223-60.
Deraniyagala: “The Pleistocene of Ceylon,” CNH S, July 20, 1958.
2 H. Stoudt: “The Physical Anthropology of Ceylon,” CMES, No. 2 (1961).
The Taxonomy of the Australoid Subspecies 425
These skulls are not, as Dr. Deraniyagala first thought ( and as I
did, too, when I saw them in Ceylon), Australoid. They are
Caucasoid with, in a few cases, a Negroid overtone. The bones
are thin, the brow ridges light, and at least on T-24-B, there is
no nuchal crest. The mandible of this last skull is lightly built and
delicate, and it has a sharply pointed chin. Like the upper jaw it
shows a marked alveolar prognathism, which occurs in 7 per cent
of living Veddas. The nose is narrow and moderately prominent,
but the entrance to the nasal passages is guttered.
My present opinion, which may have to be revised after these
skulls have been cleaned, repaired, and definitively studied, is that
Balangoda man did not differ subspecifically from the living Ved-
das. If he showed some Australoid features, this should surprise
no one because his island is located just south of a Caucasoid-
Australoid zone of contact, and these skulls are not very old.
The Taxonomy of the Australoid Subspecies
In this chapter I have demonstrated, at least to my own satis-
faction, that in southeast Asia and Indonesia a pr e-sapiens popu-
lation of the genus Homo evolved, from the very beginning of
the Middle Pleistocene onward, through three known stages into
a congeries of modern races, the Australian in sensu strictu, the
Tasmanian, the Papuo-Melanesian, and the Negrito. Even the
Negritos differ among themselves. The Andamanese of the main
archipelago lack the pronounced steatopygia of the Onges of
Little Andaman, and both kinds of Andamanese are more infan-
tile facially than the Philippine Negritos. The Tasmanians, though
possessing Negroid hair, were morphologically close to the Aus-
tralian aborigines, and perhaps even closer to the spiral-haired
Melanesians of New Caledonia. We can be reasonably sure that
the Negritos of the southeastern quadrant of the Old World be-
came small independently of the African Pygmies, but we do
not know whether one or several populations underwent dwarfing
in the area under consideration.
Among the Australians themselves regional differentiation may
be seen, and the peripheral tribes tend to be more primitive
426 Pithecanthropus and the Australoids
morphologically than those in the central desert. In New Guinea,
the Papuans are vastly differentiated regionally. Blond hair can
be seen in some of the central Australian tribes, in New Caledonia,
and in Fergusson Island, one of the D’Entrecasteaux group lying
off the southeastern tip of New Guinea. A strict application of the
taxonomic rules stated in Chapter r might give the status of sub-
species to some of the populations living in this geographical
quadrant.
But I prefer to call them local races, particularly as they have
certain features in common. All have broad noses, wide interorbi-
tal distances, dark skins; and the adult males have beards. Den-
tally they are also closely similar, particularly in the relationship
between the size of the cheek teeth and the anterior length of the
skull, as expressed by Flower’s index (see Chapter 8), in which
all Australoids are megadont, whether they are full-sized or
pygmy-sized, curly-haired or straight-heared, or whatever.
As we have noted before, stature and hair form divide the
Australoids into several groups. In Chapter 3 dwarfing was ex-
plained; the dwarfing of some of the Australoids in geologically
recent times probably followed similar patterns. Because dwarfing
shows on the skeleton, we have been able to trace it in at least
one area, Indochina. More difficult to explain is the distribution
of hair form, which is of two kinds: straight to ringlets, and
negroid. The negroid type of hair is peripheral geographically to
the straight, being found in Tasmania, Papuo-Melanesia, and
among the mainland and Indonesian Negritos. In India it also
seems to be peripheral to straight hair among the predominantly
Australoid tribes. Unless we can postulate multiple mutations
within our geographical quadrant, and can also explain why
these mutations followed a marginal geographical pattern, we
are almost obliged to consider this difference in hair form ancient.
Either the straight hair of the central Australians is the result of
some kind of selection, like the juvenile and female blondism of
the aborigines living in the central desert; or it represents influ-
ences derived from the vanguard of the first Mongoloid invasions
which reached Indonesia before the last wave of Australoids had
left for Australia; or it is the result of undetected Caucasoid
The Taxonomy of the Australoid Subspecies 427
movements from India— these are the only explanations I can
think of.
Summing up, we may divide the Australoid subspecies of man-
kind into three races, characterized as follows:
( 1 ) Full-sized, with straight or wavy hair: Australoid proper
(2) Full-sized, with negroid hair: Tasmanian and Papuo-
Melanesian
( 3 ) Pygmy-sized, with negroid hair: the Negritos
Among at least the first two races, local populations differ con-
siderably in evolutionary grade, and some of them come closest,
of any living peoples, to the erectus-sapiens threshold.
In my own opinion, looking at this chapter in retrospect, its
most important conclusion is not that the Pithecanthropus- Wad-
jak evolutionary line has been established, for this fact has been
acknowledged before, but that we now have enough skeletal ma-
terial from the period between 40,000 years ago and the present
to carry that line through to modern times. No longer need we
rely on hypothetical invasions from an unknown center.
It also occurs to me that the transition from Homo erectus to
Homo sapiens in this quadrant was caused by gene flow from a
Mongoloid source. This is suggested by, among other things, the
extraordinary facial flatness of Wadjak, and by the fact that dur-
ing the entire span of human history as we know it, the Aus-
traloids and Mongoloids were in contact, like the United States
and Canada, over an open frontier. In the following chapter we
shall try to follow the evolution of the Mongoloids from Sinan-
thropus to the present-day peoples of that area.
SINANTHROPUS
AND THE MONGOLOIDS
The Living Mongoloids and the Skeletons
of Their Ancestors
TT
nuke the three different but related races of Australoids
whose common origin we tried to trace in Chapter 9, the Mon-
goloids of the world, from Madagascar to Tierra del Fuego, are a
relatively homogenous subspecies. They have coarse, straight,
black head hair which grows very long and grays only in extreme
senility; and they rarely become bald. Neither sex has very much
body hair, and the adult male has little beard. They have a tend-
ency to facial flatness, protruding malars, widely separated and
shallow eye sockets, nasal bones which invade the frontal bone
deeply, large incisors which are usually shoveled, relatively long
bodies and short lower segments of the arms and legs, along with
small hands and feet.
Aside from these similarities, they are of all sizes above the
Pygmy, varying according to standard zoological rules with lati-
tude and altitude. Their skin color also tends to vary regionally,
but not as much as in the Caucasoid subspecies. Some of them,
like the southern Chinese, have very flat noses, whereas others,
like the Nagas of Assam and the American Plains Indians, have
aquiline ones. But these differences are minimal compared to
those found in most other subspecies, and a common origin for
all Mongoloids is clearly indicated.
Of all the living subspecies of man they are also the most dif-
ferentiated, and the least like any of the others. One can see a
MAP 10
primitive European in an Australian aborigine and an African
Negro in a Melanesian, but except for their resemblances to the
African Bushmen in facial flatness and skin color, and except for
the presence of a few flattish faces in north-central Europe, the
Mongoloids stand alone. The questions that must be answered in
this chapter are: How far back do the features that characterize
the Mongoloids go in prehistory? Can they, as Weidenreich said,
be derived from Sinanthropus? Let us look at available evidence.
This evidence, not including that from the Americas, is given
on Table 19. It covers only fourteen sites and sixty-odd individuals,
over forty of whom are Sinanthropi. Only twenty-one individuals
from thirteen sites stand between Sinanthropus and the late pre-
historic Chinese. However, these thirteen sites are widely scat-
tered from the bend of the Yellow River to the Pacific, and from
Kwangtung and Szechuan in China to Honshu in Japan. Also,
they are well distributed on the time scale. Their temporal and
430
Sinanthropus and the Mongoloids
TABLE 19
EARLY SKELETAL MATERIAL FROM
CHINA AND JAPAN
Time
Place
Material
Middle
(1) Choukoutein, original ex-
Remains of 40+ individuals
Pleistocene
cavations and those of 1959;
including 14 calvaria, 12 man-
all Mindel II, ca. 360,000
dibles, and 147 teeth; named
years old
Sinanthropus pekinensis, Pithe-
canthropus pekinensis, and Pith-
ecanthropus sinensis
( 2) Ting-tsun, Shansi, late
Middle Pleistocene
Lungtung Cave
Three teeth
( 3) Changyang, Hupei, same
Fragment of maxilla, 3 teeth
( 4) Mapa, Kwangtung, same
Fragment of calvarium
( 5) Ushikawa Quarry, Honshu,
Japan, same
Fragments of humerus
Upper
( 6) Tze-Yang, Szechuan
Skull, male, 14-15 years
Pleistocene
(7) Liu-Kiang, Kwangsi
Cranium and pelvic bones,
male, 40 years ca.
( 8) Sjara-Osso-Gol, Ordos
One left upper lateral incisor
( 9) Ti-Shao-Gou-Wan, Ordos
Fragment right parietal and
half a femur
(10) Aichi, Honshu, Japan
Fragment calvarium and os
coxae
Late Upper
(11) Choukoutien, Upper Cave *
Seven individuals of which 3
or Early Post-
skulls are described
Pleistocene
(12) Kait’o-Tung Cave, Kwangsi
Skull base, palate, teeth
Post-
(13) Chalinor, Manchuria
Two skulls
Pleistocene
(14) Yokosuka, Honshu, Japan,
Early Jomon, 6450 B.c.
One adult male skeleton
* W. C. Pei, W. P. Huang, C. L. Chiu, and H. Meng, in VP, Vol. 2, No. 4 (1958), pp. 226-9, call this
site Final Pleistocene or the so-called Postglacial of Europe, on the basis of the fauna.
spatial distributions are thus better than what we had to work
with in Chapter g.
Sinanthropus pekinensis: Time, Place, and People
Choukoutien, or Chou Gate Inn, is a limestone cliff
thirty miles south of Peking. In it are breccia-filled clefts, long
the haunt of dragon-bone collectors whose finds end up in Chi-
nese pharmacies. In 1903, in a Peking drugstore, K. A. Heberer
Sinanthropus pekinensis: Time, Place, and People 431
found a human tooth which was recognized as that of a fossil
man. From then on for twenty-four years various paleontologists
and anatomists, having traced the tooth back to Choukoutien,
worked in or watched the site, which had once been a large cave.
In 1927 Birger Bohlin, a Swedish paleontologist, found a molar
in situ which Davidson Black, professor of anatomy at Peking
University, named Sinanthropus pekinensis. From 1927 through
1937 the site was worked continuously under the direction of
W. C. Pei and Pere Teilhard de Chardin. Franz Weidenreich
joined them before the excavations were completed and described
the finds,1 except for one skull already described in print by Black.2
In the 1950 s the Chinese Communist government resumed work
at the site and a new mandible was discovered and described in
1959-3
Early in World War II, at the beginning of the Japanese occu-
pation of China, all the Sinanthropus skulls were lost in an acci-
dent or military action while being transferred from Peking to the
S. S. President Harrison. No one seems to know what happened
to them. We blamed the Japanese for their disappearance and
now the Communist Chinese blame us. All we have left is a set
of casts made in the basement of the University Museum in Phila-
delphia, a lone tooth which is in Sweden, and the new mandible
from China, in addition to a set of detailed monographs by
Weidenreich and a few other publications.
In the course of excavation, between 1927 and 1937, Pei, Teil-
hard, and their associates removed from the breccia not only the
celebrated fossil human remains but also many tools, some char-
coal, the seeds of the hagberry, which is a kind of wild cherry,
1 F. Weidenreich: “The Mandibles of Sinanthropus pekinensis,” PSNSD-7,
No. 3(1936).
: “The Dentition of Sinanthropus pekinensis,” PSNSD, No. 1 (1937).
: The Ramification of the Middle Meningeal Artery in Fossil Hominids
and Its Bearing upon Phylogenetic Problems,” PSNSD, No. 3 ( 1938 ) .
: The Extremity Bones of Sinanthropus pekinensis,” PSNSD, No. 5
(i94i).
: “The Skull of Sinanthropus pekinensis,” PSNSD, No. 10 (1943).
" D. Black: On an Adolescent Skull of Sinanthropus pekinensis . . . ” PSD,
Vol. 7, No. 1 (1927), pp. 1-28. (Black’s monograph describes Skull E, or No. 3
of Weidenreich).
3 J. K. Woo and T-K. Chao: “New Discovery of Sinanthropus Mandible from
Choukoutien,” VP, Vol. 3, No. 4 ( 1959), pp. 169-72.
432 Sinanthropus and the Mongoloids
and the broken and splintered bones of many animals. As the
human remains had been broken in the same fashion, it was clear
from the start that the Sinanthropi had been eaten by cannibals,
presumably also Sinanthropi.
With a few exceptions, like that of Rhodesian man, who
crawled into a narrow cave to die alone, nearly all the available
skeletal material from the Middle Pleistocene, and some from the
Late Pleistocene also, commemorates the ancient practice of man
eat man. This does not mean that for hundreds of thousands of
years every human being ended up in someone else’s stomach; but
if you are eaten, your bones have a better chance of being pre-
served for posterity than if your body is simply abandoned. Being
tossed into a garbage dump is better from the archaeological point
of view than being left for the wolves and hyenas on the lone
prairie.
Moreover, the great cannibals of the world are farmers whose
tiresome starch diets make them crave meat, as for example the
Caribs, Papuans, and Azande. Hunters eat one another only when
starving, not just for protein, but for calories. As the Sinanthropi
were hunters, we may assume that the scraps of well-picked hu-
man bone which they threw into the cleft at Choukoutien repre-
sented moments of extreme hunger. Had the entire population
been eaten over the thousands of years that this site was occupied,
the excavators would have found the remains of thousands of
Sinanthropi instead of a scant forty.
As the bones were both broken and scattered, it was difficult
for Weidenreich to decide which pieces belonged together, and
the correlation of mandibles to the correct crania, loose teeth to
jaws, and long bones to skulls was nearly impossible. Nevertheless,
he managed to assemble fourteen adult calvaria, which, although
fragmentary, cover between them virtually the whole cranium
except for the basal region around the foramen magnum. As
usual, this had been broken off so that the brain could be ex-
tracted.
Table 20 lists the cranial and mandibular specimens studied by
Weidenreich; calvarium 3, however, was described in 1927 by
Black. Of the 147 teeth, Weidenreich was able to study all but
two. Thirteen are milk or deciduous teeth; 134 are permanent.
Sinanthropus pekinensis: Time, Place, and People
433
TABLE 20
THE SINANTHROPUS SPECIMENS BY
SEX AND AGE
Calvaria F acial Bones Mandibles
o. Sex Age No. Bone Sex Age No. Sex Age
l-B
M
adult
2-D
?
adult
3-E
M
juvenile
4-G
M
juv. or adol.
5-H
M
adult
6-1
F
adult
7-1
M
adolescent
8-J
F?
juv., 3 yrs.
9-J
M
juv., 6 yrs.
10-L
M
adult
11-L
F
adult
12-L
M
adult
13-0
M?
adult
14-UC *
M
adult
*UC =
upper cave
1 frag, max-
ilia
M
adult
2 frag, zygo-
matic arch
M
adult
3 frag. max.
+ 5 teeth
F
adult
4 }/2 palate
F
adult
5 frag. max.
+ 6 teeth
M?
adult
6 frag. max.
+ 4 teeth
M
adult
A-2
M?
adult
B-l
F
juvenile
B-2
M
juvenile
B-3
F
juvenile
B-5
F
juvenile
B-6
F
juvenile
C-l
F
juvenile
F-l
M
juvenile
G-l
M
adult
H-l
F
adult
H-4
F
adult
M-WCf
F
adult
t WC = Woo & Chao, ’59.
Although only lower teeth are represented in the deciduous col-
lection, there is at least one of each tooth, and among the perma-
nent teeth every tooth in both jaws is accounted for. Three of the
milk teeth are called male, ten female; of the permanent teeth
fifty-five are called male, four male( ?), and five female.
The postcranial bones, which are not listed above, consist of
seven fragmentary femora designated as A-i to A-7, of which A-2
is called adult female and the others adult male; two fragments
of humerus, B-i and B-2; one piece of clavicle; and an os lunatum,
or wristbone. The last four bones are called adult male. None of
these bones could be related to the skulls or teeth with any cer-
tainty.
As he did later with Pithecanthropus and Solo, Weidenreich
sexed the Sinanthropus skulls, teeth, and long bones on the basis
of size. This was possible because they varied considerably. It
was an arbitrary procedure which he himself questioned in his
later writings. Whether or not it was justifiable is important be-
cause the living Mongoloids do not have that much sexual di-
morphism. In fact some of them have very little. If the Mongol-
oids are descended from Sinanthropus, then either sexual di-
434
Sinanthropus and the Mongoloids
morphism has decreased in the Mongoloid subspecies since that
time or Weidenreich’s sexing is incorrect.
Another possibility is that the size difference might reflect evo-
lutionary change, because the fissure in Choukoutien is very deep
and bones were recovered over a vertical range of 150 feet.
Davidson Black, who began the excavation, originally divided
this range into fifteen units of 10 feet each, designated by the
TABLE 21
LOCI AND SEX OF SINANTHROPUS
SPECIMENS
Specimens by Loci and Sex
Loci and Sex Correlated
*
Locus
Skulls
Mandibles
Dentitionsf
Loci
Males
Females
Total
A
F,M
A-C
7
5
12
B
M
F,F,F,F,M,
F
C
F
M,M
E-J
7
10
17
D
E
F
M
M
F,M
F,M,M
K-0
5
5
10
G
M
M
Total
19
20
39
H
M
F,F
F
I
F,M
F
J
M
K
F
L
F,M,M
M
M
F
M
N
F
0
F,M
* Only specimens designted as M or F and by level are included. Those designated as M?, F?, ?, and
U.C. are excluded.
f These are sets of teeth which Weidenreich assembled.
letters A to S, and called loci. Locus A was at the top and Locus O
at the bottom. We have twenty-seven definitely sexed skulls
and mandibles from these loci, as indicated on Table 21 and
plotted there in the form of a correlation table. There is no sta-
tistically significant difference between the loci, as combined
above, and Weidenreich’s sex designations,4 and therefore no evi-
dence that the differences in skull size and jaw size which
Weidenreich interpreted as sex differences were really evolution-
ary in origin.
However, Weidenreich’s vindication must be tempered by an-
4 As determined by the chi-square method: P = 33.
Sinanthropus pekinensis: Time , Place, and People 435
other factor, the nature of the deposit itself. The fill, which later
became brecciated, was only partly deposited by occupation.
Some of it fell into the cave from above as debris. Although the
breccia is stratified, one cannot be sure which bone entered the
cave from in front and which fell in through the hole in the roof,
from the land surface above. Bones and other hard objects falling
100 feet or more do not necessarily stay where they land, but can
tumble and roll about. If they hit a sloping floor they can come to
rest at some distance from the point of contact. So we cannot be
completely sure that Weidenreich was right after all.
Recent work on the geology of the fissure indicates that the
deposits were laid down over an extensive period.5 They consist
of three gravels: basal, lower, and upper. The Basal Gravels
were formed, apparently during the Giinz glaciation, partly as a
result of gravels and red clay being washed into a large cave
from outside, and partly by internal deposition. The Lower
Gravels were formed by river water depositing various pebbles,
gravels, sands, and clays. Their age is stated to be First, or Cro-
merian, Interglacial. The Upper Gravels were formed by internal
deposition, in what was still a large cave, in Mindel-Elster time.
Before this, the cave was not habitable. The Sinanthropus remains
are, according to this new work, all of Second Glacial, or Mindel-
Elster, age, but this part of the Pleistocene lasted tens of thou-
sands of years. There was still time enough for a little evolution-
ary change within a single local population.
The new Chinese geological findings have been supported by
Kurten’s equally new paleontological determinations. He calls the
date of the site Mindel-Elster II, about 360,000 years ago accord-
ing to the argon-potassium dating method,6 later than the time of
the Djetis Pithecanthropi, and perhaps contemporary with Trinil.
It was the time when Crocuta crocuta, the living spotted hyena,
which had recently evolved in India, was crowding its older
W. P. Huang: Restudy of the CKT Sinanthropus Deposits,” VP Vol. a
No. 1 (i960), pp. 45-6.
Huang: On the Age of Basal Gravel of CKT Sinanthropus Site, of the ‘Upper
Gravel and the ‘Lower Gravel’ of the CKT Region,” VP, Vol. 4 No 1 (1060)
pp. 47-8.
11 B. Kurten: “New Evidence on the Age of Peking Man,” VP, Vol. 3, No 4
(1959), pp. 173-5-
436 Sinanthropus and the Mongoloids
cousin, Hyaena breviostus, out of its ecological niches in Africa,
Europe, and China. Commonest among the animal bones were
those of deer, showing that venison was a far more popular en-
tree on the menu than roast Sinanthropus.
Pollen analysis conducted in Finland on a piece of Choukoutien
breccia gives the following percentages for tree pollens: 33 per
cent pine, 28 per cent beech, 9 per cent alder, 4 per cent spruce,
3 per cent linden, 1 per cent yew, 1 per cent willow, and 1 per
cent sea buckthorn. Of the nonarboreal pollens, 11 per cent be-
longed to grasses, 4 per cent each to sedges and the rose family,
3 per cent to the sagebrush- worm wood tribe ( Artemesia ), and
1 per cent each to the goosefoots ( Chenopodeaciae ) and crow-
berry (Empetrum) .
As the pollen-bearing breccia sample was collected by the Sino-
Swedish expedition before the excavation had reached below the
first few loci, it probably represents the end rather than the be-
ginning of the Sinanthropus occupation. At that time the climate
was cooler than it is today. Choukoutien then lay near the border
zone which separates the northern coniferous belt, or boreal
forest, from the temperate steppe. Its hills were clad with pine and
spruce, their slopes bearing berries which grow today in the very
north of Maine. What is left of the local vegetation at present,
after thousands of years of intensive use of the land by man, is
typical of steppe or parkland, and Choukoutien now lies nearer
the edge of the evergreen tropical forest of southern China than
that of the northern forest of Manchuria.
Sinanthropus had fire. His implements were good enough for
working skins crudely. He must have been clever enough to keep
from freezing during the winter months, and he was very likely
physiologically adapted to cold, at least to the extent of the mod-
ern Alakalufs. One wonders how long he had lived in China be-
fore the span of the Choukoutien site. Many stone implements
are turning up in China nowadays, but none is much cruder than
his, except in the south. Did he make his entry with the spotted
hyena, or did he evolve locally from some even more primitive
kind of man? We do not know, but at the rate Pleistocene studies
are moving in China, before long we may.
The Taxonomy of Sinanthropus
437
The Taxonomy of Sinanthropus
A s previously stated, in 1927 Davidson Black created the name
Sinanthropus pekinensis to describe one tooth. This term was im-
mediately applied to other remains excavated in Choukoutien as
they appeared, and is still in general use. Yet Boule and Vallois
renamed Sinanthropus in 1952, calling him Pithecanthropus pe-
kinensis, and in 1957 Piveteau dubbed him P. sinensis.8 In recent
years P. pekinensis has been widely accepted in Britain and on
the continent. As I agree with Simpson, Mayr, Washburn, and
others in calling all fossil men Homo, Sinanthropus will be called
Sinanthropus without italics, and the other names given him will
be mentioned again only in the index.
Still, because the new name. Pithecanthropus pekinensis, has
been accepted by many prominent scholars, it may be worth
while to trace the origin of this term. It seems to have come from
a remark in 1943 by Weidenreich: “Sinanthropus differs from
Pithecanthropus in characters which have not so much phyloge-
netic as racial bearing.” 9 In other words, they belong to more
or less the same grade but to different lines.
Two years later G. G. Simpson, in his authoritative opus on
mammalian taxonomy, had this to say: “All specimens of fossil
hominids that differ in any discernible way from Homo sapiens,
and some that do not, have at one time or another been placed
in different genera. Almost none of these anthropological ‘genera’
has any zoological reason for being. All known hominids, recent
and fossil, could well be placed in Homo. At most, i Pithecanthro-
pus (with which f Sinanthropus is clearly synonomous by zoologi-
cal criteria) and f Eoanthropus (if the apelike jaw belongs to it)
may be given separate generic rank. Perhaps it would be better
for the zoological taxonomist to set apart the family Hominidae
7M. M. Boule and H. V. Vallois: Les Hommes Fossiles (Paris: Masson et Cie-
1952), p. 145-
8J. Piveteau: Traite de Paleontologie (Paris: Masson et Cie; 1957) Vol 7
( P. sinensis is used in captions of illustrations. )
9 Weidenreich: “The Skull of Sinanthropus pekinensis,” PS-NS-D Vol 10
ws No. 127 (1943). ’ ' ’
438 Sinanthropus and the Mongoloids
and to exclude its nomenclature and classification from his stud-
• 99 1
les.
It is hard to see how either of these statements warrants the
definitive pooling of Pithecanthropus and Sinanthropus in a ge-
nus of their own apart from Homo, except as a tentative taxonomic
device which, by i960, had lost its usefulness.
The Sinanthropus Brain Case
We have five Sinanthropus brain cases (see Table 37) com-
pared to three for Pithecanthropus and six for Solo. No. 3 is a
juvenile and No. 2 has no breadth or height measurements. As all
had been opened for extraction of the brain, none has a complete
base; but No. 12 still had the posterior margin of the foramen
magnum and in No. 11 enough pieces were in contact with each
other to permit its restoration.
Omitting the juvenile No. 3, the cranial capacity of the re-
maining four ranges from 1,015 to X225 cc ., almost exactly the
same as the Solo range, and a notch above that of Pithecanthro-
pus. Sexual dimorphism in the cranial capacity of Sinanthropus
can neither be established nor disproved because only one
measurable skull out of four is called female.
In comparison with the Solo skulls, those of Sinanthropus are
both shorter and narrower, but of about equal height. How, then,
can they have the same capacities? Table 22 holds the answer. In
their internal dimensions the Sinanthropus skulls are 14 mm.
longer than the Solo average and 7 mm. narrower, and both sets
are of about equal height. In a long narrow brain, apparently, two
millimeters of length equal one of breadth ( 14 = 7 X 2), provided
that the third dimension, brain height, remains constant.
The difference between the two groups in external skull length
is due to the more massive growth of brow ridges and nuchal
crests in Solo, particularly noticeable in No. 5. His length of 219.5
mm. is excessive in the genus Homo as a whole, and is a record
for fossil men anywhere, yet his internal length of 175 mm.
1 G. G. Simpson: “The Principles of Classification and a Classification of
Mammals,” BAMN, Vol. 85 (1945), p. 188.
439
The Sinanthropus Brain Case
TABLE 22
INTERNAL DIMENSIONS OF THE
SINANTHROPUS AND SOLO SKULLS
Sinanthropus Solo
Aver- Aver-
Skull No.
8
10
11
12
age
1
5
6
9
10
11 age
Age
juv.
ad.
ad.
ad.
ad.
ad.
ad.
ad.
ad.
ad.
Sex
M
M
F
M
F
M
F
?
F
M
Length
156
184
166
175
175
161
175
153 * 164 *
159*
157 * 161
Breadth
122
128"
' 128
129
128
130
138
129
137
138
129 135
Height
105
102
110
106
103 ’
" 108
109*
‘ 105*
100*
109 * 107
* Qualified by (?) or circa.
matches that of Sinanthropus 12, whose total external length is
only 195-5 mm-
Two points emerge from this comparison. ( 1 ) Although identi-
cal in size, the two sets of brains were different in shape. (2) Al-
though he lived at least 200,000 years before Solo man, Sinanthro-
pus was already more highly evolved than the latter in the reduc-
tion of bony superstructure about the brain case, just as he had a
larger brain than his own probable contemporaries, the Pithe-
canthropi of Trinil. No better evidence could be found, consider-
ing the paucity of skulls at our disposal, to show that in different
anthropogeographical regions evolution proceeded at different
rates during the Pleistocene.
The paths followed in the evolution of the human brain can
best be traced by studying the interiors of the skulls of persons
of different ages, particularly the youthful and very young. In the
Sinanthropus collection, No. 3, who died at the age of eight or
nine, is available for study. The internal length of his brain was
20 mm. short of the figure for his elders; its breadth was within
6 mm. of theirs, and he equalled them in brain height. Judged by
these figures, the gross shape of the Sinanthropus brain was mod-
ern at the age of eight, and it lengthened and flattened out later.
It would be fine if we could say that the brain of No. 3 by the
age of eight or nine had not lost its infantile condition of having
a bent floor, but we cannot because the appropriate bones are
missing. However, much of the temporal bone is present, and the
sapiens type of structure which includes torsion leaves its mark
in this region. In the area around the internal ear, No. 3 alone
440
Sinanthropus and the Mongoloids
of all the Sinanthropi could pass for H. sapiens, which leads us
to suggest that the sapiens condition in both brain proportions
and brain floor anatomy is neotenous.2 This evidence indicates that
the difference in brain form, as apart from brain size, which dis-
tinguishes H. sapiens from II. erectus was a product of one or
more neotenous mutations.
It is possible that H. modjokertensis, the ancient companion of
Pithecanthropus 4, who died while his fontanelle was still open,
also anticipated modern man in brain anatomy as it certainly did
in gross proportions.
The configuration of what is left of the basal portion of the
occipital bone after brain-picking suggests this. In any case, we
have evidence that both the Pithecanthropus-Solo and Sinanthro-
pus populations and successions of populations bore within them
the genetic capacity for evolutionary change into H. sapiens.
Whether they made this change by themselves or were aided by
an injection of genes from other populations is a different ques-
tion.
Another result of a close examination of the inside surfaces of
the skulls of the Pithecanthropus, Solo, and Sinanthropus adults
is that all three had a wide separation between the frontal and
temporal lobes. Each of them has a pronounced internal ridge or
Sylvian crest which is rudimentary or absent in fossil European
skulls and modern crania, as well as in ape skulls. It is an early
human trait and not a pongid heritage. Its absence in H. sapiens
is therefore again neotenously human rather than gerontomorphi-
cally apelike.
2 So important is this point that I quote Weidenreich in full, for the benefit
of those with sufficient anatomical knowledge to understand the terminology.
In discussing the internal surface of the temporal bone, Weidenreich said: “As to
other features of the posterior surface the only noteworthy difference between
Sinanthropus and modern man concerns the apertura externa aqueductus vestibuli.
In the latter this slit opens into the impressio cerebellaris and the part covering
the slit projects more or less forming thereby the anterior boundary of the im-
pressio. In Sinanthropus Skull V . . . and all the other adult specimens ( right
and left side) the cover of the aperture appears as a distinct eminence and the
slit opens into a recess situated beneath the eminence. Only the juvenile Skull III
shows conditions similar to modern man. The porus acusticus internus (pai), the
fossa subarcuata, and the apetura externa canaliculi cochleae (acc) are like such
structures of modern man. Weidenreich: “The Skull of Sinanthropus pekinensis,”
PS-NS-D, Vol. 10 (1943), p. 68.
The Sinanthropus Brain Case 441
These skulls also have sagittal crests inside the frontal bone,
separating the frontal lobes of the two cerebral hemispheres. This
crest serves as an extra, unseen buttress to brace the skull. It is
never found in apes and is present, but smaller, in living men.
Actually, it is part of the archaic human architectural system of
passing the stress of chewing up the mid-line of the face rather
than to either side, as among apes and most if not all Australo-
pithecines. The size of the crest depends on the amount of stress
and on the angle between the vertical plane of the face and the
slope of the forehead. When the facial profile and profile of the
forehead form a straight line, as in apes and some modern peo-
ple, the sagittal axis of the frontal bone is directly in the path of
stress and internal bracing is unneccessary. The facial-frontal angle
in Sinanthropus was close enough to 450 to make such a strut use-
ful.
The inner surfaces of the Sinanthropus brain cases also show the
paths of the middle meningeal arteries, which feed the outer
sheath of the brain.3 In fossil man and the Australopithecines these
arteries split into two branches below the point at which their
imprints are seen in the parietal region. Once they appear they
take one of two patterns, as follows.
(A) The anterior branch crosses the Sylvian fissure and sub-
divides into an anterior or bregmatic branch, and a median or
obelionic branch. The posterior or lower temporal branch starts
out equal in size to the anterior branch and extends over the
lower rear border of the parietal bone, but it forks little.
(B) The anterior branch is larger than the posterior, and
both have more and finer fingers. Between them they cover the
surface of the brain which they feed more closely and finely, and
they are not as widely separated, so that they give the appearance
of being a single system.
Type A is characteristic of the Australopithecines, Pithecan-
thropus, Sinanthropus, and apparently of Solo, although in the
latter skulls the imprints are less easily read than in the others. It
is also found in a piece of temporal bone of Early Middle Pleis-
tocene date found at Ternefine in Algeria, and in the Late Upper
3F. C. Howell: “European and Northwest African Middle Pleistocene Homi-
nids,” CA, Vol. 1, No. 3 (i960), pp. 195-232.
442 Sinanthropus and the Mongoloids
Pleistocene skull from Broken Hill, Northern Rhodesia. Type B is
characteristic of the earliest skulls from Europe, which come from
the Second or Great Interglacial, and of modern men in general.
Seen from the outside, the Sinanthropus skulls are long and
low, with their greatest breadths directly over the mastoids. Like
the Pithecanthropus crania they suggest a poorly raised loaf.
However, the frontal region is quite different. Instead of a grad-
ual slope upward and backward from glabella, the frontal bone
rises steeply to form a brow ridge, sweeps back horizontally, then
bends and rises abruptly for a short distance and curves back-
ward to bregma, the point where the frontal bone joins the parie-
tals in the sagittal line. Bregma itself is situated directly over
porion, the top of the earhole; in modem skulls it lies forward of
this position. In some Sinanthropus skulls the frontal torus is solid,
in others it contains a small frontal sinus. In Pithecanthopus and
Solo it encloses large frontal sinuses.
Behind the brow ridges lies a deep postorbital constriction,
comparable to that of the Pithecanthropi but not of Solo. In this
part of the frontal bone the area of temporal muscle attachment
invades the lower part of the forehead as it does in modern Mon-
goloids. The average distance between the temporal crests for
Sinanthropus is 94.7 mm. compared to Solo’s 108.7 mm. No two
sets of skulls could differ from each other more, in this feature,
than these.4
The lambdoid region is flattish, the occiput bun-shaped, and
the zygomatic arches were undoubtedly flaring. Beyond their
rearward anchor on the temporal bone they continue over the
earhole as a supramastoid crest, but do not move on to join the
nuchal crest. Although this bony framework of the skull is massive
and impressive, it does not equal that of the early members of the
Pithecanthopus line, as our comparison of inside and outside
skull diameters indicated. As far as we can tell, the foramen
magnum was set farther back in the skull base in Sinanthropus
than in the Pithecanthropus-Solo skulls. The mastoid processes
4 Skull 12 has two sets of temporal crests, an inner and an outer. I found the
same anomaly on an Alakaluf Indian hi Chile. The muscles do not extend beyond
the outer crests.
1
444
Sinanthropus and the Mongoloids
C
TZE-YANG FEMALE
Fig. 6o Profiles: from Sinanthropus to the Upper Cave Male. A. Weiden-
reich’s restoration of a Sinanthropus female; B. The Mapa skullcap (after Woo,
1959); C. The Tze-Yang skull, female (after Woo, 1958); D. The Upper Cave
male. No. 101 (after Weidenreich, 1943). During the late 1950’s Chinese paleon-
tologists found several specimens which bridged the gap in the Mongoloid sequence
between Sinanthropus ( Homo erectus) and the Upper Cave people ( Homo
sapiens), thus establishing the fact that Mongoloids evolved from one species to
the other in East Asia. Although the Mapa skullcap is too fragmentary for a certain
diagnosis, at least on present evidence, the Tze-Yang skull, from the Early Upper
Pleistocene, is definitely sapiens.
of Sinanthropus are well developed, but instead of descending
vertically, as one would expect, they point inward.
The Sinanthropus brain cases also possess a number of pe-
culiarities of a racial rather than evolutionary nature. All but
No. 3 have Inca bones — supernumerary bones separated from the
occipital bone just below lambda, which is the point where the
two parietals and the occipital meet in an inverted Y. They also
have exostoses, or bony growths on the edge of the auditory
meatus, or bony earhole. In No. 3 a fissure in the tympanic plate,
known as the infantile gap, remains open, whereas in European
skulls it closes well before the age of five. In living Mongoloids it
tends to remain open into adulthood, having thus been found, as
well, in an incidence amounting to 33.3 per cent of skulls in some
The Face of Sinanthropus 445
American Indian series, 32 per cent of prehistoric skulls from
Guam, and 12 per cent of more recent North Chinese crania.
The Face of Sinanthropus
The upper facial bones of Sinanthropus are represented by
the six pieces listed in Table 20 and by the nasal bones and por-
tions of orbits forming part of the brain cases. These can best be
studied from Weidenreich’s reconstruction of a female skull,
which includes a mandible. Some of the dimensions of the recon-
struction may be seen on Table 23.
TABLE 23
FACIAL DIMENSIONS OF SINANTHROPUS
(FEMALE) AND WADJAK 1
Sinanth.
Sinanth.
female t
Wadjak 1
female t
Wadjak 1
Minimum Frontal
84 mm.
99 mm.
Facial Index
79.7
87.1*
Bizygomatic
148
140(7)
Upper Facial
52.1
52.9
Index
Biorbital
111
115
Fronto-parietal
64.9
66.9
Index
Interorbital
25
29
Cranio-facial
105.7
94.6(7)
Index
Bicondylar
124
140*
Zygo-frontal
64.7
70.7
Index
Bigonial
103
110 *
Zygo-gonial
69.6
79.6
Index
Total Face Height
118
122*
Nasal Index
57.2
56.0
Upper Face Height
77
74
Orbital Index
81.9
78.6
Nasal Height
52.5
50
Palatal Index
75.1
70.0(7)
Nasal Breadth
30
28
Orbital Height
36
33
Orbital Breadth
44
42
Palate Length
52
60
Palate Breadth
39
43
* The Wadjak 1 cranium and the Wadjak 2 mandible were combined.
t The Sinanthropus female is Weidenreich’s reconstruction. No Sinanthropus skull of either sex was
whole enough to permit these measurements.
It is not a particularly large face. All of the dimensions given
could be matched in living populations. Compared with Pithecan-
thropus 4, it belongs in a different order of magnitude, and it is
even smaller in eight of fifteen dimensions than Wadjak I when
446 Sinanthropus and the Mongoloids
the latter is equipped with the W-2 mandible, which may be a lit-
tle big for it. The importance of this comparison is that it shows
the people of northern China 360,000 years ago to have had
faces, at least in the female sex, of modern size, and smaller in
many dimensions than the Australoids of Java who lived at least
300,000 years later.
Both the upper face and the nose are relatively long, like those
of the modern Chinese; and despite the heavy brow ridges, the
Sinanthropus orbits have modern Mongoloid dimensions, al-
though they are less rounded than in the recent skulls. From the
SINANTHROPUS MAXILLA No. 0-1 A MODERN NORTH CHINESE SKULL
Fig. 61 Alveolar Procnathism in Sinanthropus and
in Modern Chinese. (Drawings after Weidenreich,
1937- )
modern point of view the only excessive dimensions of the Sinan-
thropus face are the bizygomatic diameter and the palate length.
These reflect evolutionary status more than racial affiliation.
The nasal bones are wide both at top and bottom, and flat,
meeting the zygomatic (or malar) bones 5 in a gently rounded
fashion rather than forming an angle as in the Causasoids. At the
top they meet the frontal, which they invade deeply, in the
form of a T, whereas in the Caucasoid face the nasofrontal suture
either curves upward or looks like the head of an arrow | . The
nasal opening is broad, as it is among all early fossil men, and
the lower border is guttered as in modern Mongoloids. In profile
the face is very prognathous, most of the prognathism being pro-
duced by the outward and downward curvature of the maxillae
5 This bone is officially known as os zygomaticum, but is also popularly known by
its old name, malar, which I shall continue to use.
447
The Mandibles of Sinanthropus
and palate, although some of it comes from the extension of this
curvature upward to the nasal region. In modern Chinese skulls
an almost equal amount of alveolar prognathism is occasionally
found.
Although the brow ridges make them look square, the orbits
are not low. They fall within the modern Mongoloid size range;
and if the brow ridges were reduced or removed, the similarity
would be more apparent. The floor of each orbit opens directly
and simply from the malar without lipping or sill, and the outer
sides of the orbits are relatively straight, whereas in Caucasoids
and Australoids this edge curves to the rear, giving the orbit a cut-
away appearance and making the nasal skeleton appear more
prominent. In this detail Sinanthropus resembles not only modern
Mongoloids but also the orang, the gorilla, and the Australopithe-
cines.
In Sinanthropus, as in modern Mongoloids, the temporal mus-
cle was attached farther forward than in the Pithecanthropus-
Australoid skulls, and it pushed the malar bone forward and also
invaded the frontal bone over and behind the brow ridges; the
temporal lines are much closer together than in Sinanthropus’s
grade-mate, Solo. Were one to enlarge the Sinanthropus brain by
about 300 cc., reduce the brow ridges, shorten the palate, and
reef in the zygomatic arches by about 15 mm., it would be hard
to tell this specimen from that of a modern Mongoloid of one kind
or another, at least in the upper part of the face, which is the
racially critical “mask” area.
The Mandibles of Sinanthropus
From the lips downward the similarity still exists, but it is
heavily camouflaged by the fact that the Sinanthropus mandibles
are morphologically primitive and fit the evolutionary grade to
which their great antiquity entitles them. The laboratory designa-
tions, sexes, and ages of the twelve which have been studied are
given on Table 20.
Only three are complete enough for detailed study: the adult
male G-i, the adult female H-i, and the juvenile female B-i (see
Fig. 62). H-i was used with cranium No. 11 by Weidenreich in
44§ Sinanthropus and the Mongoloids
Fig. 62 Mandibles; Sinanthropus and Ternefine 3. From the side: A. Sinan-
thropus G-i, male; B. Sinanthropus H-i, female; C. Ternefine 3, male. From
above: D. Sinanthropus G-i, male; E. Sinanthropus H-i, female; F. Ternefine 3,
male. Weidenreich found that the Sinanthropus mandibles were of two sizes. The
larger ones he called male, and the smaller female. Both are stout, prognathous,
and large-toothed. Both have multiple mental foramina. The Ternefine mandi-
bles from Algeria are also large and robust and resemble the Sinanthropus speci-
mens closely in many details, but Ternefine 3 has an extremely sloping ascending
ramus and a very high coracoid process. The latter feature is also found in
Australopithecines. Seen from above, the Sinanthropus mandibles spread far apart
behind the dental arc, so that the condyles are widely separated. This is also a
modern Mongoloid feature, and in it Ternefine 3 resembles Sinanthropus and the
modern Mongoloids. (Drawings C and F after Arambourg, 1955; all others after
Weidenreich, 1943.)
449
The Mandibles of Sinanthropus
his reconstruction of a complete female skull (see Table 23).
The new female mandible described by Woo and Chao closely
resembles H-i but is much less complete.
In eight measurements the female mandible H-i attains 85.9
per cent of the size of the male G-i. In living Mongoloid popula-
tions from China, Korea, and Japan, the figure is 92.4 per cent.
This difference in sexual dimorphism between ancient and living
Mongoloids is of the same order of magnitude as that found in
the cranial series.
The Sinanthropus mandibles as a group fall into the size range
of other fossil men of the same general time span, including
Javanese, European, and North African specimens. All are about
equally chinless, except that in the female H-i the chin line is
angular rather than curved. The angle of inclination, which is the
angle between the tooth line of the jaw and the slope of the
chin line (see Fig. 62 and Table 24) comes to 63° in the Sinan-
thropus mandibles, including the 1959 specimen. In comparison,
the angles for Pithecanthropus B, Heidelberg (Sinanthropus’s
European contemporary), and Wadjak 2 are 58°, 63°, and 65°.
In modern Australian aborigines it averages 75° and in living
Mongoloids and Europeans about 90°.
In evaluating the relative thickness of the mandible in fossil and
recent jaws, anthropologists use an index of robusticity, which is
thickness of the bone times 100 divided by the height of the
mandibular body. The two measurements, thickness and height,
are taken at the same point on the jaw when possible, but this
point may differ because some jaws are fragmentary. A usual
location is at the symphysis, on the mid-line of the jaw between
the two lower median incisors. In Pithecanthropus B the symphy-
sis is missing. Therefore a substitute index has been calculated at
a point between the median and lateral incisors, which is present
in this specimen. This index comes to 52.5, which means that the
thickness is 52.5 per cent of the height. In three Sinanthropus jaws
the figures are 36.6, 38.9, and 46.4, indicating a much less massive
bone near the chin line. However, when the same index is taken
at the conventional spot, which is the level of the mental foramen
(a hole in the bone usually situated under the point where the
second lower premolar and first lower molar meet ) , the figure for
450
Sinanthropus and the Mongoloids
TABLE 24
ANGLES OF INCLINATION AND INDICES OF
R0BUSTICIT7 OF SINANTHROPUS AND
OTHER MANDIBLES f
Angle of Inclination
APes: Men:
Orang
44°
Pithecanthropus B *
58°(?)
Gorilla
47°
Sinanthropus (6)
61° (59.0-63.5)
Chimpanzee
50°
Heidelberg
63°
Ternefine (3)
65° (62-70)
Australopitheeines :
Modern Australians
75°
Modern Whites
90° ca.
Robustus
58°
Choukoutien Upper
91°
Meganthropus
58°
Cave
Index of Robusticity (at Mental Foramen Level)
Apes:
Men:
Orang
50.8
Sinanthropus (4)
57.1 (48.3-62.3)
Gorilla
50.8
Ternefine (2)
55.5 (?) (52?-59?)
Chimpanzee
49.5
Pithecanthropus (2)
55.4 (47.2-63.6)
Heidelberg
48.8
Australopitheeines :
Modern Australians
45.6
Wadjak 2
42.9
Robustus *
60.0 (?)
Modern Means
Meganthropus
58.4
(non-Australian)
38-42
* Reconstructed from drawings or photographs,
t For further details, see Table 38 in Appendix.
the four Sinanthropus mandibles comes to 55.4, against a mere
44-5 f°r Pithecanthropus B. Thus a racial difference is evident:
the Pithecanthropus mandible is the stouter in front, and those
of the Sinanthropi more robust in mid-branch.
In Sinanthropus and in other early fossil men, the gonial angle
is blunted so that it becomes two angles of about 450 each. The
area between the two corners is the seat of attachment of the
masseter muscles on the outside of the bone, and of the internal
pterygoid muscle on the inside. Because both these muscles are
concerned with the rotary motion of the jaw in chewing, this
similarity merely reflects the common habit of heavy chewing
in unrelated lines. In Sinanthropus, as in the Eskimos and other
northern Mongoloids, the gonial angles are bent outward, making
the lower part of the face look very wide. This again is partly a
function of chewing.
The ascending rami, which connect the tooth-bearing body of
the mandible to the skull, are of moderate height in Sinanthro-
45i
The Mandibles of Sinanthropus
pus, as in other early fossil men. As we saw in Chapter 7, the
ascending rami of the Australopithecines are very long and high.
This means only that the faces of the various lines of Homo erectus
were diagnostically shorter than those of the Australopithecines
yet discovered. Also the Sinanthropus mandibles, like those of
other Homines erecti, have genial tubercles projecting from the
insides of their symphyses, like most modern men and unlike
most, if not all, Australopithecines.
While conforming to the general Homo erectus pattern in the
details mentioned above, Sinanthropus had two special peculiari-
ties, the torus mandibularis and multiple mental foramina. The
Fig. 63 Tohus Mandibularis. Cross-
section through the exaggerated torus
mandibularis of the mandible of a pre-
historic Chinese, in line with the first
lower premolar. ( Drawing after Wei-
denreich, 1936.)
FIRST PREMOLAR
torus mandibularis is a swelling of the jawbone on the tongue
side, concentrated between the level of the canine and that of the
first molar. This is solid, heavy bone, with no spongy interior.
All the Sinanthropus jaws have it, and no other early fossil man
had it. The torus mandibularis is found among some 15 per cent
of the contemporary Chinese, whereas among Eskimo popula-
tions it varies in frequency between 42 and 97 per cent. It is
found, as well, in 68 per cent of mediaeval Icelandic jaws. Neo-
lithic Japanese, Lapp, Ostiak, prehistoric Chinese, Ainu, and
prehistoric Scandinavian jaws all have it in ratios varying from
62 to 17 per cent; and it is just as frequent among northern
Caucasoid jaws as it is among those of northern Mongoloids. If
this bone is an adaptation to heavy chewing it is difficult to under-
stand why others, particularly the Neanderthals, lacked it, unless
they had some other adaptation to the same function. Oddly
452 Sinanthropus and the Mongoloids
enough, it is also found in an extinct deer which the Sinanthropus
folk ate.c We are reminded of other cases of convergence in special
environments — for example, the case of prehensile tails in differ-
ent kinds of South American mammals, including primates. As
with the prehensile tail, torus mandibularis is hereditary.7
The other peculiarity, multiple mental foramina, is even more
mysterious. In every jaw in which either or both sides of the bone
lying immediately under the second lower premolar and first
lower molar is preserved, more than one foramen may be seen.
There are seven such jaws and the number of holes ranges from
two to five. In other jaws, new and old, there is usually only one
such perforation through which the blood vessels and nerves that
service the lower part of the cheeks pass out of the body of the
bone. None of the other early fossil man jaws has this anomaly,
except for Heidelberg and the Ternefine mandibles from Algeria,
which have two. The latter resemble the Sinanthropus jaws so
closely in other respects that the two sets of jaws could have come
from a single population. Later on, multiple mental foramina turn
up in some of the European Neanderthals. In any case these two
features, torus mandibularis and multiple mental foramina, indi-
cate as well as anything could the extreme genetic isolation of the
Sianthropus population.
The Teeth of Sinanthropus
Thanks to Weidenreieh, we have more information on the
teeth of Sinanthropus than on those of any other fossil human
population.8 His series consists of 147 teeth, 83 of which are still
in their original positions in jaws and 64 of which are loose. Of the
147 teeth, 134 are permanent, including 52 uppers and 82 lowers.
Thirteen are deciduous, all lowers. The total is said to represent
about 32 individuals, 20 adults and adolescents with permanent
teeth, and 12 juveniles with milk teeth. This is as large a sample
as we get from some living populations.
6H. D. Kahlke: “On the Evolution of Pacliyostosis in Jaw Bones of CKT Giant
Deer, Megaceros Pachyosteous (Young),” VP, Vol. 2, No. 3 (1958), pp. 117-34.
7 M. Suzuki and T. Sakai: “A Familial Study of Torus Palatinus and Torus
Mandibularis,” AJPA, Vol. 18, No. 4 (i960), pp. 263-72.
8 Weidenreieh: “The Dentition of Sinanthropus pekinensis.”
The Teeth of Sinanthropus 453
Like the skulls and mandibles the teeth are of two sizes, rela-
tively large and relatively small. Large ones are found in two
large mandibles and one large “male” skull, and small ones are
found in two small mandibles and one small “female” skull. If we
agree with Weidenreich that the large teeth are male and the
small ones are female, then we have the permanent teeth of ten
male adults and adolescents and of six male juveniles, and also
of ten adult-adolescent and six juvenile females.
Table 39 gives the length and breadth of the Sinanthropus
permanent teeth and comparable dimensions for Pithecanthropus
from the two available specimens, No. 4 and mandible B, as well
as the crown dimensions of the Upper Cave specimens, those of a
series of modern Chinese males, and the modern length and
breadth ranges for all races.
The Sinanthropus teeth are large in both mesiodistal length
and labiolingual breadth diameters, but all except two of the
sixteen kinds of teeth (uppers and lowers, incisors, canines, etc.)
fall within the ranges of living populations. One exception is the
upper canine. In five of the six specimens studied, the mesiodistal
length exceeds the modern range; and one of them, with a length
of 10.5 mm., is the largest such tooth in the world. The other
exception is the second upper premolar. Two of twelve of these
exceed the modern range in the labiolingual breadth.
In general Pithecanthropus had much larger teeth than Sinan-
thropus, with seven of twelve ( all sixteen are not represented in
Pithecanthropus) exceeding the Sinanthropus range. The two
populations also differ in the ratio between the combined mesio-
distal length of the three upper molars and the total length of the
entire upper tooth row, from the upper median incisor to the
upper third molar. In Sinanthropus the molar length is only 41.5
per cent of the total row length; in Pithecanthropus 4 it is 45.2
per cent. These differences are retained, more or less, in modern
Mongoloid and Australoid peoples.
In Sinanthropus the upper first molar is the largest of the three
and the upper third the smallest. This is the usual sequence in
modern populations. In Pithecanthropus 4 the upper second molar
is the largest and the first and third are nearly equal in size. In the
lower jaw the second molar is the largest in both Sinanthropus
454
Sinanthropus and the Mongoloids
and Pithecanthropus B, but in Sinanthropus the first molar is
much larger than the third, while in Pithecanthropus B the first
and third molars are nearly equal in size. These differences be-
tween the Javanese and Chinese Homines erecti in tooth size re-
flect the differences already noted in the stoutness of the lower
jawbones of the two races at different locations — that in Sinan-
A
B
c
D
Fig. 64 The Continuity of Mongoloid Teeth: Shovel Incisors from Sinan-
thropus to the Recent Chinese. Upper Median Incisors: A. Sinanthropus;
B. Ting-tsun; C. Upper Cave; D. Recent North Chinese. Upper Lateral Incisors:
E. Sinanthropus; F. Ting-tsun; G. Sjara-Osse-Gol; H. Recent North Chinese. This
sequence of upper and lower incisors shows a continuity of tooth form in the heart
of the Mongoloid realm from the Middle Pleistocene to the present. ( Drawings B
and F after Movius, 1956; all others after Weidenreich, 1937. )
thropus the emphasis is on the front of the mouth whereas in
Pithecanthropus it is on the molars. The whole facial structure of
Sinanthropus, as previously noted, is concentrated on its forward
portion, which is still true of Mongoloids.
The relationship between the teeth of Sinanthropus and those
of living Mongoloids is shown more convincingly in morpho-
455
The Teeth of Sinanthropus
logical characteristics than in gross dimensions. Of these the most
conspicuous is the peculiar shovel-like shape of the upper incisors,
described in Chapter 8. All five of the upper median and both of
the upper lateral incisors are shoveled. Not only are the edges of
the teeth raised, but they are actually wrapped around on the
lingual side, and they have one to three fingerlike tubercles or
ridges running down the lingual surface from its upper inner
rim half way to the cutting edge. Incisors comparable to those
of Sinanthropus have been found in the earlier South African
Australopithecines and in some of the fossil jaws from North Af-
rica, and the element of lingual ridging occurs in the incisors of
living great apes.
The upper canines of Sinanthropus are large and long-rooted,
and their crowns project beyond the level of the incisors and
premolars. Instead of points, as in apes and modern men, the
lower canines have cutting edges. The upper canines are shoveled
and fingered, braced and ornamented, in the manner of the upper
incisors. Starting with the canines and moving backward, all the
teeth have cingulums, or collars, sometimes completely circular,
sometimes incomplete, just above the necks of the teeth, at the
bases of the crowns. This feature is characteristic of apes and is
slightly developed in some of the Australopithecine teeth. In
Pithecanthropus it is present but not pronounced, and it is not
characteristic of modern man of any race. Like Pithecanthropus,
the orang, and some of the Australopithecines, Sinanthropus had
fine wrinkles on the grinding surfaces of his molars. Both the
cingulum and the wrinkling are more characteristic of an early
grade of human dental development than of a particular line.
All the permanent molars and premolars of Sinanthropus, and
all the milk molars, are taurodont; that is, each tooth has an en-
larged pulp cavity extending downward into fused roots, as de-
scribed in Chapter 8. A taurodont tooth can be worn down much
lower than an ordinary tooth. This condition, absent in Pithe-
canthropus, is not unique with Sinanthropus, but is found in a
number of populations, ancient and modern, particularly among
the Middle Pleistocene people of North Africa, the European
Neanderthals, and living Eskimos, American Indians, and Bush-
men.
456 Sinanthropus and the Mongoloids
Taurodontism seems to have a selective advantage when the
workload of the teeth is too great for their surface area, as, for
instance, in a cold climate, where the teeth are used for softening
skins (Neanderthals and Eskimo), or as when, after the teeth
have been reduced by dwarfing, the capacity for heavy chewing
is still needed (Bushmen). In Sinanthropus, skin dressing was
probably the critical factor. Taurodontism is hereditary. All of the
peoples who have it or had it ( with the possible exception of the
Neanderthals, to be studied in the next chapter) are in one way
or another members of either the Mongoloid or the Capoid racial
line of descent.
The Leg Bones of Sinanthropus
Weidenreich has described seven fragmentary femora,
numbered 1 to 7. Numbers 4 and 5 are believed to be a pair.
Not one has a complete head or distal end. All except No. 2,
which is smaller than the others, are called male. Only No. 1 and
No. 4 are complete enough to permit reconstruction of their total
lengths, 400 mm. and 407 mm; these indicate a stature of about
five foot one and a half inches, or 156 cm, similar to that of living
Japanese, Eskimo, and Ainu, and shorter than that of Pithecan-
thropus.
Several peculiarities set these femora apart from those of most
modern men. The walls of the shaft are extraordinarily thick and
the medullary canal occupies only 35 per cent of the total diame-
ter (33 per cent transversely and 38 per cent sagittally) as com-
pared with about 48 per cent for other ancient men of later pe-
riods. The Pithecanthropus femora were also of normal thick-
ness.9 Among the apes only the orang has thick femoral walls.
In Sinanthropus the dense bone of the walls runs far up on
the neck of the femur where modern femora contain mainly
spongy matter. Furthermore, the trajectorial grid system of the
less extensive spongy part is composed of exceptionally coarse
9 Judging from the photographs of the broken shafts of Nos. 2 and 3 in the
Leyden collection (Weidenreich, 1941, plate 29) and a photograph of No. 1
which shows a break on the inside of the lower shaft (R. Martin, 1928, p. 1153).
457
The Upper Extremity of Sinanthropus
fibers which are laid down in a different pattern from that seen in
modern femora.
The Sinanthropus femora are also extremely flat, fore-and-aft,
like those of apes; this feature is also found in some other popu-
lations, including Neolithic Chinese and modern Fuegians.
Uniquely, however, the Sinanthropus femora have only vestigial
pilasters ( bony struts along the center of the backs of the shafts ) ;
the linea aspera ( a ridge superimposed on the pilaster ) , which is
found in all human specimens, consequently lies flat on the surface
on the bone. Although the shaft is bent no more than in modern
specimens, the peak of the curve lies near the knee-end instead
of in the middle as in other femora, including those of Pithecan-
thropus.
The Upper Extremity of Sinanthropus
There is only one piece of humerus, and both ends of it are
missing. This bone was probably about 324 mm. long, or 30 mm.
longer than those of Japanese and Ainu, who have the same femur
length. If Weidenreich’s reconstruction of it is correct, either it
belonged to a taller individual than the owners of the six femora,
or else he had proportionately longer arms. Like the femora, the
humerus is extremely thick-walled, although its general form is
slender. Otherwise it is completely modern. It has, however, an
extreme development of the tubercle for the attachment of the
deltoid muscle, the function of which is to raise the arm side-
wise. Similar tubercles have been found on the humeri of Neo-
lithic Chinese and modern Fuegians. The Fuegians are great
paddlers; what Sinanthropus and the Neolithic Chinese did to de-
velop such powerful deltoids is unknown.
We have also one slightly defective clavicle in the Sinanthropus
collection. As this is an extremely variable bone in modern man,
it is not surprising that the specimen at hand has no special
features; it is slender, highly curved, and heavily ridged for the
attachment of powerful muscles, including the deltoid.
There is also one os lunatum, a wrist bone, which is also com-
pletely human, although it is small; but so, apparently, are those
of many Mongoloids.
458
Sinanthropus and the Mongoloids
The Position of Sinanthropus in the Human Family Tree
Even these scanty observations make it evident that the
postcranial skeleton of Sinanthropus differed from that of Pithe-
canthropus, and that both differed from the curiously assorted
skeletons of Australopithecus. Among these animals the head of
the femur was not rotated as far forward as in man, and the distal
condyles of that bone had peculiarities of their own. In the
Zinjanthropus leg bones the fibula is thicker, compared to the
thickness of the tibia, than in man as a genus, and this suggests
that Zinjanthropus walked on the outside of his foot. Pithecan-
thropus had long, slender, modern-style femora, and his successor,
Solo man, a modern tibia. Sinanthropus, however, had short,
heavy leg bones, with peculiar bone webbing in the head of the
femur, narrow marrow cavities, a relatively short tibia, and much
bowing of both bones.
Sinanthropus differed from Pithecanthropus in many other
anatomical details. His brain case was larger, as large as Solo’s.
In some details of cranial anatomy, particularly in the configura-
tion of the inner surface of the temporal bone, he was more ape-
like than Pithecanthropus. His frontal bone had a constriction
behind the brow ridges, whereas Pithecanthropus’s brow rose in
a smoother, more gradual slope. Pithecanthropus (at least Num-
ber 4) had the largest palate, and their teeth were as different
as teeth could be within a single species.
These comparisons suggest that the relationship between Pithe-
canthropus and Sinanthropus, which most physical anthropolo-
gists believe in, was not a close one. It also suggests that either
several related hominids acquired the erect posture independently
or, by the time Sinanthropus came along, different lines had been
as rigorously differentiated by the evolutionary process from the
neck down as from the neck up. The currently popular nostrum
that except for their skins all men are alike from the neck down
is nonsense.
Sinanthropus was a peculiar type of human being who had
more features in common with living Mongoloids, regardless of
The Position of Sinanthropus in the Human Family Tree 459
grade, than with any other living subspecies. Among these com-
mon features are the following:
( 1 ) A sagittal keeling of the skull vault, found among Eski-
mos and North Chinese. Pithecanthropus also had this, as do
modern Australians and Tasmanians.
(2) Inca bones, found in three or four of five skulls. These
are found in 15 per cent of the American Indians and are more
frequent among Mongoloids than in other races.
(3) Broad nasal bones that show little or no difference be-
tween upper and middle breadths.
( 4 ) A gently rounded contour of the nasal saddle.
( 5 ) The profile angle of the roof of the nasal passages equals
89°, a little higher than in Mongoloids, who have the highest
such angle among living men.
(6) The outer border of the orbit is set forward as in Aus-
tralopithecines, gorillas, and orangs, and the forward part of the
temporal muscle is extended anteriorly above the edge of the
brow ridge, compressing the lateral half of the orbit.
(7) The infraorbital margin is rounded and even with the
floor of the orbit, as in modern Mongols.
(8) Buccal exostoses (bony growths) of the mandible are
found in all three upper jaws of Sinanthropus; these growths are
found in from two to five per cent of the Aleuts, the Japanese, the
Lapps, and the natives of Siberia.
(9) Exostoses of the internal auditory meatus (tube of the
earhole ) .
(10) A general thickening of the tympanic plate. This and the
preceding are found chiefly among Eskimos, American Indians,
and Icelanders.
( 11 ) The “infantile gap” in the tympanic bone.
( 12 ) A special external growth on the border of the tympanic
plate, found in Sinanthropus Skull X and in no other fossil homi-
nids; it occurs in 18 to 20 per cent of the Polynesians, 12 to 30 per
cent of the American Indians, and only rarely in Caucasoids.
( 13 ) The mandibular torus.
( 14 ) Shovel incisors .
( 15) Extreme flattening of the femur, accompanied by a flat
linea aspera and a distal position of the shaft curve.
460 Sinanthropus and the Mongoloids
(16) A strongly developed deltoid tuberosity of the humerus.
( 17 ) A small wrist bone.
Weidenreich’s list, given above with a few modifications,1 im-
plies a genetic continuity with the modern Mongoloids of Asia,
Oceania, and America; but it has not been widely accepted, for
two reasons. First, some of the features which appear in other
populations that dwell in cold regions may have been acquired
adaptively and convergently. Second, most human anatomists are
reluctant to admit that more than one line of human beings could
have passed the evolutionary threshold that separates Sinanthro-
pus from the living Mongoloids. In my opinion the first objection
is more valid than the second.
Still a third objection, which has held back a number of open-
minded scientists who are willing to overlook the first two, is the
lack of skeletal material to fill the time gap between Sinanthropus
and historical races of Mongoloids. Owing to new discoveries in
China and Japan, that gap is being filled.
Late Middle Pleistocene Finds in China and Japan
Since 1954 four different finds of fossil man, made in the Far
East, have been assigned to the later part of the Middle Pleisto-
cene by their discoverers, three from China and one from Japan.
They are listed on Table 19.
The Ting-tsun Teeth
I n 1954 fourteen paleolithic sites were excavated in the region of
Ting-tsun, Shansi, northern China.2 Although they are stated to
be Middle Pleistocene by the Chinese discoverers, Movius be-
lieves that they are of Third Interglacial age. Over two thousand
1 Weidenreich: “The Skull of Sinanthropus pekinensis,” pp. 252-4.
2 H. L. Movius, Jr.: “New Paleolithic Sites near Ting T’sun on the Fen River,
Shansi Province, North China,” Quaternaria, No. 3 (1956), pp. 13-26.
T-K. Cheng: Archaeology in China, Vol. I, Prehistoric China (Cambridge:
W. Heffer and Sons; 1959), pp. 25-6.
G. Bushnell and C. McBumey: “New World Origins Seen from the Old World,”
Antiquity, Vol. 33, No. 130 ( 1959), pp. 93-101.
The Changyang Maxilla 461
artifacts found here are said to indicate an evolutionary progres-
sion based on the stone-tool industry of Choukoutien, with or
without influences from the Western world.3
The human remains from these excavations consist of three
teeth, two upper incisors and a lower second molar. They are
smaller than the Sinanthropus teeth and within the modern Chi-
nese range.4 Both incisors are shoveled in the exaggerated Sinan-
thropus fashion. The lower molar, which has five cusps and an
incipient sixth, resembles those of Sinanthropus morphologically.
Whether these teeth belonged to the Late Middle or Early
Upper Pleistocene is less important than the fact that they form a
bridge between Sinanthropus and modern Mongoloids in asso-
ciation with a stone-tool industry derived from that of Choukou-
tien. This is the continuity that Weidenreich sought and died too
soon to see.
The Changyang Maxilla 5
Three years later, a piece of fossil human maxilla, contain-
ing an upper first premolar and an upper first molar, and also an
isolated lower second premolar, were found in a cave called
Lungtung, at Hsiachungchiawan village, in the Ichang lime-
stone area 28 miles southwest of the city of Changyang in Hupei
Province. This is mountainous country; the cave is about 4,400
feet above sea level.
Although there were no artifacts in this site, faunal remains
were abundant. They belonged to the so-called Ailuropus-
Stegodon fauna ( Ailuropus is the giant panda), which was also
found with Sinanthropus at Choukoutien. Chia, who described it,
considers the fauna of this site to be of Late Middle Pleistocene
date, and to my knowledge no one has yet challenged this al-
location.
3 Movius states that they show no Western influence; Bushnell and McBurney
that they do.
4 Scale measurements of the illustrations give the upper median incisor a length
of 9 mm., the upper lateral incisor a length of 7 mm., and the lower second molar
a length of 11 mm., and a breadth of 11 mm.
5 L-P. Chia: “Notes on the Human and Other Mammalian Remains from
Changyang, Hupei,” VP, Vol. 1, No. 3 ( 1957), pp. 252-7.
462 Sinanthropus and the Mongoloids
The maxillary fragment is part of the left side only, including
the roof of the palate and the sagittal line from the nasal spine to
the tooth line; in other words, half of the upper jaw. The palate is
ribbed, as it should be; the anterior nasal spine is poorly devel-
oped and pointing forward; the nasal opening is wide, and its
lateral wall less curved than in most modern men (this is ap-
parently a primitive feature noted by the author of the mono-
graph).
The relief of the bony surface of the maxilla which covers the
root of the canine is high, indicating that that tooth, which is
missing, had a long, thick root as in Sinanthropus. However, the
three teeth that are present are intermediate in size between the
teeth of Sinanthropus and the teeth of modern Chinese, although
the root of the lower second premolar is longer than either of the
two corresponding Sinanthropus teeth.6 The enamel of these teeth
is highly wrinkled.
Morphologically and metrically the Changyang specimens
closely resemble Sinanthropus, but seem to indicate a step for-
ward in the direction of modern Mongoloid man. Because the
brain case is missing, we have too little evidence to indicate
whether or not Changyang man had crossed the threshold from
Homo erectus to Homo sapiens; but if he had not, he was well on
the way.
The Specimen from Mapa, Kwangtung
A year after the Changyang discovery, in 1958, farmers dig-
ging fertilizer in a cave in the so-called Lion Hill at Mapa, Shao-
quan Municipality (formerly Chukiang District) of Kwangtung
(Canton) Province, found a fragmentary human skull in the
midst of many other mammalian bones, including Stegodon, an
extinct elephant. The fauna indicates a Late Middle or Early
Upper Pleistocene date.7 The human specimen, already men-
6 The crown dimensions of the three teeth are: upper first premolar, length =
7.4 mm., breadth = 10.6 mm.; upper first molar, 1. = 10.8 mm., br. = 12.8 mm.;
lower second premolar, 1. = 8.3 mm., br. = 10.6 mm. The root length of the
lower second premolar is 20.5 mm., those of the Sinanthropus specimens 17.3 mm.
and 19.2 mm.
7 Woo: Fossil Human Skull of Early Paleanthropic Stage Found at Mapa,
Shaoquan, Kwangtung Province, VP, Vol. 3, No. 4 (1959), pp. 176-82.
The Specimen from Mapa, Kwangtung 463
tioned in Chapter 9, consists of the frontal bone, both parietals,
the nasal bones, and the lower border of the right orbit. It is
heavily fossilized; the sutures are all fused; it is certainly adult,
and probably male.
With this skull two questions arise at once: Is it erectus or
sapiens P Is it Mongoloid or Australoid, if either? Unfortunately,
not enough of it is left to help us answer either question com-
pletely.
The frontal bone is longer than the parietals, and bregma is
located farther back than in most modern skulls. These are both
hallmarks of Homo erectus. In the indices of the arcs and chords
of the frontal and parietal bones ( see Chapter 8 ) the Mapa cra-
nial fragment falls into the ranges of the Sinanthropus and Solo
skulls, the Rhodesian and Saldanha specimens from South Africa,
and also the European Neanderthals.8 But the absolute measure-
ments of the bones slightly exceed the Sinanthropus range, and
they lie just inside the Solo range.
Woo has drawn the skull on a simulated eye-ear plane (see Fig.
60) and drawn a line from glabella, which is present, to
opisthion (the rearmost point of the occiput), which is postulated.
The height of the skull above this line (82 mm.) is the same as
that of Sinanthropus 10, the largest of the Sinanthropi. As the
Mapa skull is only 7 mm. thick at bregma, compared to 8.8 mm.
for Sinanthropus, and as the skull appears to have been more
rounded, its capacity probably exceeded that of Sinanthropus 10
and the largest of the Solo skulls ( 1,225 cc)> and fell easily within
the modern range.
Morphologically it is again intermediate, both in grade and in
line. The brow ridges are heavy, and shaped like those of Solo
rather than those of Sinanthropus. But the frontal bone behind
the ridges is markedly constricted as in Sinanthropus; and the
nicking of the frontal profile just above glabella is moderate, and
intermediate. The nasal bones are wide and the nasofrontal suture
almost straight. The frontal sinus is larger than in either Sinan-
thropus or Solo, and extends laterally over the eye sockets. The
orbital borders are rounded, not square as in Sinanthropus; but
8 The Mapa frontal arc = 134 mm., chord = a 18 mm., index = 88.0. The
figures for the parietal bone are 114 mm., 110 mm., and 96.5.
464 Sinanthropus and the Mongoloids
they are also rounded in modern Mongoloids. The orbits them-
selves are high, as in Mongoloids. Also the lower border of the
orbit projects forward, as in Sinanthropus and the Mongoloids.
Woo believes that the Mapa skull had evolved to the same
grade as the European Neanderthals, to what he called the Early
Paleanthropic stage. In view of the history of the Neanderthals,
which will be recounted in the next chapter, I agree with him in
this diagnosis but not in the comparison. The Mapa skull seems to
stand at the threshold between the two grades of Homo. If it was
not sapiens, it was very close to being so. In any case, it represents
a higher stage of human evolution than Sinanthropus himself,
which is the most important conclusion we can reach. As to its
race, it seems to me to be mostly if not entirely Mongoloid; and in
ways in which it differs from Sinanthropus, as in the shape of the
orbits, it is a link between Sinanthropus and the modern Mon-
goloid peoples.
The Humerus Shaft from Ushikawa Quarry, Japan
Until after the Emperor’s official declaration that he was
no longer to be considered divine, the search for fossil man in
Japan was not vigorously pressed. During the 1950’s, however, it
got off to a late but profitable start. In 1957, a laborer working in a
limestone quarry in the Ushikawa district, five miles from the city
of Toyohashi in Aichi Prefecture, east-central Honshu, found a
number of bones at a depth of 70 feet. Among these bones were
two broken pieces of the shaft of a human humerus. The fauna
with which they were associated is called Late Middle Pleisto-
cene. No artifacts were found. These humeral fragments, which
are from one bone because the two sections fit together, have been
described by H. Suzuki and F. Takai.9
The piece is 70 cm. long, and comes from almost exactly the
middle of the shaft, which is believed to have had a total length of
230 mm. Suzuki, who thinks it female, compares it to the humeri
of nineteenth-century Japanese women, the mean of which is
265.5 mm. This suggests a stature for the Ushikawa woman of
9 H. Suzuki and F. Takai: “Entdeckung eines Pleistozanen Hominiden Hu-
merus in Zentral-Japan,” AAnz., Vol. 23, No. 2/3 ( 1959), pp. 224-35.
The Upper Pleistocene Woman from Tze-Yang, Szechuan 465
only 135 cm., or 4 feet 5 inches, which would not be very tall for a
Pygmy woman.
A detailed study of the various diameters and circumferences
taken at different loci on the shaft show that it differs considera-
bly from those of the modern Japanese, being, among other
things, narrower at the proximal end and thicker at the distal.
Moreover, the walls of the shaft are very thick compared to the
width of the marrow cavity; the walls comprise 55 per cent of the
diameter, compared to 40 per cent for recent Japanese. In this
respect it resembles the limb bones of Sinanthropus. This is a most
unusual bone, and although it has been described exhaustively,
we must, along with Suzuki and Takai, await further discoveries
before it can be properly evaluated.
The Upper Pleistocene Woman from Tze-Yang, Szechuan 1
Turning again to Table 19, we find five items from the Far
East labeled as Upper Pleistocene. The first is a skull from west-
central China, from the mountainous province of Szechuan, which
today is inhabited not only by Chinese but also by Tibetans,
Lolos, and other non-Chinese-speaking tribesmen. The skull was
found in 1951 by railway workers in a bank of the Huangshanchi
River, Tze-Yang District, Szechuan. Associated with it was a
rather crudely made bone awl and an extensive fauna.
The animal bones, although mixed together in the deposit,
could be separated into two lots on the basis of color, degree of
fossilization, and fluorine content. The older fauna includes
Stegodon orientalis, Rusa unicolor, and Rhinoceros sinensis, Steg-
odon being an extinct elephant, and Rusa simply Cervus rusa,
the extinct deer found at Choukoutien. This fauna belongs to the
Middle Pleistocene. The younger fauna includes Muntiacus (the
muntjak deer), Mammonteus primigenius ( Mammuthus , accord-
ing to Simpson, a mammoth), and Homo sapiens, this being the
Tze-Yang woman herself. This fauna is Upper Pleistocene. The
sorting out of these two faunas by W. C. Pei may help clarify the
dating of other sites, particularly in south China, where the faunas
are also scrambled.
1 W-C. Pei and Woo: “Tzeyang Paleolithic Man,” IVPM, No. 1, 1957.
466 Sinanthropus and the Mongoloids
The skull, identified as a female over fifty years old, consists of a
nearly complete vault, with both parietals intact, all of the frontal
bone except for the internal part over the orbits, all of the occipital
bone except for the piece immediately behind the foramen mag-
num, the left temporal, the left great wing of the sphenoid except-
ing the base, and small pieces of nasal bone adhering to the
frontal. Separate and without point of contact with the vault is
the palate, including parts of the maxillae and the lower part of
the nasal opening. All the upper teeth are gone except for the
broken-off root of the left upper second premolar. The right upper
second premolar and all three left molars had been lost before
death; the others had fallen out after death. There is evidence that
the woman had suffered a serious dental disease.
The cranial measurements given on Table 37 indicate that the
skull was quite small but well within the female range of both
Metal Age prehistoric and recent North Chinese series. Com-
pared to Sinanthropus 11, her skull is short, narrow, and high, and
more voluminous by 200 cc. The minimum frontal, measured on
the photograph, was probably narrower than either Sinanthropus
11, as reconstructed by Weidenreich, or the modern female mean.
The nasal opening breadth, also reconstructed, was narrower than
that of Sinanthropus 11 and close to the modern figures. The pal-
ate dimensions are modern, and the teeth were probably also of
modem dimensions.
Although the Tze-Yang woman was essentially sapiens, her
skull shows several archaic features. In the endocranial cast the
cerebral fossae of the occipital bone are wider and deeper than
the cerebellar fossae. This condition is reminiscent of Sinanthro-
pus and the Neanderthals, and indicates that the cerebellum had
neither expanded nor been pushed down by the cerebral hemis-
pheres to the extent seen in most modern skulls. Also, on the inner
surface of the parietals the impressions of the middle meningeal
artery are archaic in pattern. The anterior ramus, although the
larger, has fewer branches; the posterior ramus is the more intri-
cately branched.
The outside of the skull shows heavier brow ridges than usual
for Mongoloids, a relatively long frontal bone with bregma placed
to the rear of its modern situation, a rounded occiput, and a swell-
The Upper Pleistocene Man of Liu-Kiang, Kwangsi 467
ing above the mastoids. The squamous portion of the left temporal
bone is smaller than in most modern skulls of the same size. Also,
according to Pei and Woo, “in modern man the zygomatic process
of the temporal bone and its backward extension of the supra-
mastoid crest lie nearly parallel to the eye-ear plane. In Sinan-
thropus it forms an acute angle of 30 0 with the eye-ear plane,
while in Tze-Yang man it forms an angle of about 200.” 2
This excellent and detailed study, the high points of which have
been given here, makes it evident that the Tze-Yang skull is an
early sapiens form retaining some Sinanthropic features combined
with, for the most part, modern proportions. The position of the
temporal attachment on the frontal and its general morphology,
including that of the lower nasal aperture, also indicate that it
was Mongoloid, the only atypical feature being that the nasofron-
tal suture is slightly rounded instead of running straight across.
This is not enough to upset Pei’s and Woo’s racial diagnosis.
The Upper Pleistocene Man of Liu-Kiang, Kwangsi
Kwangsi is the next province to the west after Kwangtung,
where the Mapa specimen was found. With Yunnan, it is the gate-
way through which Mongoloids crept down the fingerlike ridges
that form the steep watersheds between the Irrawaddy, Salween,
Mekong, and Red rivers, into the steaming jungles of southeast
Asia, to replace the Australoids and Negritos who had evolved
there.
In a cave called Tungtienyen, 10 miles southwest of Liuchow in
central Kwangsi, workmen found, in 1958, an almost complete
human skull.3 No artifacts were with it, but it was accompanied
by many animal bones of the familiar stegodon-giant panda
fauna. Although it was heavily fossilized, the skull was found in
red soil, whereas most of the deposit was yellow. This fact led
Woo to conclude that its date is Upper Pleistocene rather than
Middle Pleistocene, as would have been indicated had it been
embedded in the yellow material.
The cranium is nearly complete, but there is no mandible. Also
2 Pei and Woo: op. cit., p. 40.
3 Woo: “Human Fossils Found in Liukiang, Kwangsi, China,” VP, Vol. 3,
No. 3 (1959), pp. 109-18.
L
468 Sinanthropus and the Mongoloids
found were four thoracic and five lumbar vertebrae; five pieces of
rib; a sacrum; a right ilium-ischium combination, but no pubic
bone; and two pieces of femur, one from each leg. All but the
femora are said to have belonged to a male about forty years old.
The leg-bones may have been his, or they may have been part of
a female. They are of a different color from the other bones.
As the figures given on Table 37 indicate, it is a large and
capacious skull, fully modern in the dimensions of its brain case;
but its face is low, its nose short and wide, and its orbits low. Its
palate is of moderate size, and its teeth the same. The incisors
were shoveled— at least the lateral ones were. Although the one
remaining median incisor was too worn to tell, the median incisors
are always more shoveled than the lateral ones if this trait is pres-
ent. Curiously enough, this man lived to be over forty without
cutting his upper third molars. This, too, is a Mongoloid trait.
The brow ridges are a little heavy for modem Chinese, but not
for peripheral Mongoloids like some American Indians. The posi-
tion of bregma is still too far back for a modern skull, as the frontal
part of the sagittal arc is more than one third of the total. The rear
profile of the skull shows a moderate amount of lambdoid flatten-
ing, present in both Sinanthropus and the European Neander-
thals. We have no information on the inside surface of the skull.
Is this skull, then, Mongoloid or Australoid? Woo believes that
it is Mongoloid, of a primitive type, and points out that low faces
and low orbits were common elsewhere in Upper Pleistocene
times, particularly in Europe; this was a phase through which
skulls of different lines passed independently. The form of the
nose, with its guttered rim; the slight alveolar prognathism; the
shape of the temporal lines on the forehead; and the teeth are all
Mongoloid. The molars are prominent in the forward plane, as
they should be. The skull’s frontal index of facial flatness, simotic
index (which indicates the degree of lateral curvature of the nasal
bones), and rhinial index (which expresses the degree of flatness
of the mid-face) are all within the ranges of modern Mongoloid
peoples, and the first two are within the Sinanthropus range. The
rhinial index of Sinanthropus cannot be calculated.4
4 For the Liu-Kiang skull, F.I.F.F. = 15.7; Simotic I. = 28.3; Rhinial I. =
32.4. For Sinanthropus 12, F.I.F.F. = 16.1 (?), Simotic I. = 30.0.
The Liu-Kiang Postcranial Bones 469
Woo’s conclusion that Liu-Kiang man was a Mongoloid form of
Homo sapiens still in process of evolution seems correct, except
that the skull deviates somewhat from the Mongoloid line in an
Australoid direction, as one would expect from an ancient skull
from southeast China, the contact zone between the Mongoloid
and Australoid peoples.
The Liu-Kiang Postcranial Bones
If all we had from Liu-Kiang was the skull just described,
our problem would be simple, but we have a number of post-
cranial bones, listed on Table ig — and this raises complications.
Four thoracic vertebrae, numbers g, 10, 11, and 12, which are the
bottom four, have a combined ventral body height of 84.0 mm.,
which is short for living peoples; and the five lumbar vertebrae
have a combined height of ng.i mm., which is even shorter. Of
the living peoples occupying eastern Asia only the Sakai are
known to approximate these figures, and the Sakai are aboriginal
hunters, of the Malay Peninsula, of unknown origin, whose physi-
cal appearance is largely Australoid. The Sakai are not Pygmies,
but they are very nearly so.
The accompanying sacrum is somewhat flattish and small, with
a length of gg.2 mm., a breadth of 86.5 mm., and a length-breadth
index of g3.8. It is too small for Mongoloids or for Australian
aborigines, and falls into the size range of the Andamanese Negri-
tos. It is nearly triangular, tapering toward the distal end, very
much like a Sakai sacrum illustrated in Martin’s Lehrhuch der
Anthropologies The piece of pelvic bone, which is an ilium and
ischium without the pubic bones, matches the sacrum, and the
acetabulum is rotated somewhat forward, as in Mongoloid pelves.
The leg bones consist of two broken pieces of shaft, without
condyles and of unequal length. Probably the total femur length
was about 370 mm, and the stature calculated from the femur
length about four feet eleven inches ( 150 cm.) if a male or four
feet nine inches ( 145 cm. ) if a female. These figures are on the
5 1928 edition, Vol. 2, p. 1085.
470
Sinanthropus and the Mongoloids
upper border of the Pygmy range, but they are consistent with
the size of the vertebrae and pelvic bones.
Despite their shortness, the femurs are stout, with sagittal and
transverse diameters close to those of Sinanthropus, and the de-
gree of flattening of the shafts is intermediate between that of
Sinanthropus and that of the modern Chinese.*' Also, the marrow
canal occupies 37.8 per cent of the shaft diameter, at its nar-
rowest point, in the Liu-Kiang femurs; in Sinanthropus it oc-
cupies 35.7 per cent; and in the modern Chinese 45 per cent and
higher.
One may place these three specimens and sets of specimens in
the following order of magnitude: (1) the skull is large; (2) the
femora are fairly small but rugged; (3) the trunk bones, particu-
larly the sacrum, are very small. As there is no duplication of
parts, we cannot be sure that more than one individual is rep-
resented. For a racial diagnosis each part must be taken sepa-
rately. The skull is mostly but not wholly Mongoloid, with some
features reminiscent of or adumbrating the Australoid or Negrito.
The pelvic and vertebral skeleton suggests the modern Sakai, who
are themselves an enigmatic people, and the femora suggest a
small Mongoloid. Let us hope that more material will turn up from
Kwangsi so that this mystery may be solved.
The Tooth of Sjara-Osso-Gol, Ordos
In the bed of the desert river Sjara-Osso-Gol, in the Ordos
country between the Great Wall and the bend of the Yellow River,
the late Pere Teilhard de Chardin and E. Licent found, in 1922,
one upper left lateral incisor tooth, 7.1 mm. in mesiodistal diam-
eter, which is smaller than the smallest upper lateral incisor in
the Sinanthropus series. Morphologically, however, it fits the Si-
nanthropus pattern, with heavy shoveling and a basal tubercle
projecting downward on the lingual side. It was associated with
6 Sagittal diameter = 26.2 mm.; transverse diameter = 22.0 mm. The Sinanthro-
pus figures are 28.3 mm. and 24.4 mm. The Index of platymeria (shaft-flattening)
is 67.8 for Sinanthropus, 73.7 for Liu-Kiang, and 80.2 for modem Chinese.
47i
The Remains from Ti-Shao-Gou-W an, Ordos
an Upper Pleistocene fauna and many blade implements believed
to represent the Upper Paleolithic culture of that region.7
The Remains from Ti-Shao-Gou-W an, Ordos
In another Ordos site, near the village of Ti-Shao-Gou-
Wan, two more pieces of Ordos man turned up in 1957, and they
have been given the same date as the previously described tooth.8
They are a broken parietal bone and the lower half of a femur.
The parietal bone is of modern size and shape, with a sagittal arc
of 125 mm., a chord of 110 mm., and a curvature index of 88.
These three figures are close to the modern mean for all races.
Yet the bone is a little thicker than the modern mean,9 and the
tracks of the middle meningeal artery are simple, with the poste-
rior branch larger than the anterior.
The femur half is 203 mm. long, suggesting a stature of about
five feet five inches ( 167 cm. ) if a man, or five feet three inches
( 160 cm. ) if a woman. Woo favors the latter sexing. The walls are
thick, with the marrow canal one third the total.
These two specimens, plus the tooth from Sjara-Osso-Gol, seem
to provide a continuity from Sinanthropus into the Upper Pleisto-
cene in Inner Mongolia as well as in other parts of China.
The Upper Pleistocene Remains from Central Honshu, Japan
In September 1958, six pieces of human skeleton were un-
earthed in a lens of clay in a limestone quarry in the prefecture of
Aichi, town of Mikkabi, Tadaki District. Although no artifacts
7 E. Licent, P. Teilhard de Chardin, and D. Black: “On a Presumably Pleisto-
cene Human Tooth from the Sjara Osso Gol ( South Eastern Ordos ) Deposits,”
BGSC, Vol. 5, No. 4 (1927), p. 287.
Also Weidenreich: “The Dentition of Sinanthropus pekinensis,” PS-NS-D,
Vol. 1 ( 1937), p. 21 and plate 3; and Cheng: op. cit., pp. 32-4.
8 Woo: “Fossil Human Parietal Bone and Femur from Ordos, Inner Mongolia,”
VP, Vol. 2, No. 4 (1958), pp. 208-12.
9 The thickness of this parietal near bregma is 6.5 mm.; the modem mean 5.5
mm.; and the mean for Sinanthropus 8.8 mm., with a range of 7.0 mm. to
10.0 mm.
472 Sinanthropus and the Mongoloids
were found, faunal remains were abundant, including tiger, some
kind of elephant, deer, boar, and badger. F. Tayaki of Tokyo
University has labeled this fauna Upper Pleistocene, and
H. Suzuki, who described the Ushikawa humerus shaft, is working,
at the time of writing, on the human material. These consist of
five pieces of skullcap and one fragmentary pelvic bone. The skull
pieces are two parietal fragments, two fragments of frontal in-
cluding the orbital margins, and one piece of occipital. Only the
pieces of the parietals fit together. Suzuki, in a preliminary state-
ment to the press in June 1960, said that the skull represents the
same stage of development as Cro-Magnon in Europe. For further
information we must await the publication of his final study.
The People of the Upper Cave of Choukoutien
At this point we have exhausted the human skeletal ma-
terial from China and Japan which most authorities agree is of
Upper Pleistocene date. In eastern Asia this stretch of about
150,000 years cannot be broken down as finely into subperiods as
it has been in Europe. Some of the five specimens or sets of speci-
mens that we have studied may be older than others by as much
as 100,000 or more years. Yet most if not all of them show some
racial likeness to Sinanthropus, in the skull, face, and leg bones,
although those in south China also reflect the proximity of Aus-
traloids. We have yet to discuss two lots of material which may be
dated toward the very end of the Upper Pleistocene, correspond-
ing to the final Wiirm in Europe, or even early postglacial time.
The first of these consists of the famous Old Man of the Upper
Cave of Choukoutien and his equally celebrated two wives, or,
more properly, female fellow victims.1
The Upper Cave in which they were found is a dissolution
cavity in the limestone, one which was not open in Sinanthro-
pus’s day. It contained an industry of an evolved type derived
1Pei: “A Preliminary Report on the Late Paleolithic Cave of Chou Kou Tien,
BGSC, Vol. 13, No. 3 ( 1934), pp. 327-58.
Weidenreich: “On the Earliest Representatives of Modem Mankind Recovered
on the Soil of East Asia,” PNHB, Vol. 13, Part 3 (1938-9), pp. 161-74. Also re-
ferred to in his 1943 monograph.
The People of the Upper Cave of Choukoutien 473
from the old complex of earlier days of choppers and chopping-
tools and flakes; there is no evidence in it of diffusion of European
or other Upper Paleolithic techniques. Archaeologically, the cul-
ture of these people was a local evolutionary product. The cave
soil was also crammed with fossil animal bones, including those of
hares, bears, hyenas, tigers, Sika deer, roe deer, and even ostriches
and cheetahs. Of these, the hyenas, bears, and ostriches represent
species extinct in China. Without doubt, C-14 and pollen analysis
will one day pin down the age of this deposit. Meanwhile, let us
place it in the neighborhood of about 10,000 b.c., with a wide
margin of error.
Parts of the skeletons of at least seven persons were found, but
only three skulls have been described: a man about sixty years
old, a young woman who had not yet cut her wisdom teeth, and a
somewhat older woman whose dental crowns had been worn flat.
They are numbered 101, 102, and 103. The man was given a
stature of five feet eight and a half inches (174 cm.); female No.
102 of five feet two and a half inches ( 159 cm. ) . We have no fig-
ure for No. 103. The femora of No. 101 had the same shaft-
medullary cavity ratio as those of modern north Chinese.
These people had been killed in a mass murder and left where
they lay. They had apparently not been eaten. No. 101 was killed
by an arrow or small-headed spear that pierced his skull at the
point where the frontoparietal suture crosses the temporal lines.
It was not mutilated, and was complete when found. Females
Nos. 102 and 103 suffered a less unanticipated and more horrible
death. Someone held their heads sidewise on a stone, while
someone else dropped another stone on them, squashing them so
that the bones sprang out, increasing the head height at the ex-
pense of head breadth. Furthermore, No. 102 had suffered a small
degree of cranial deformation before death, because across her
forehead stretches a furrow of the kind made by carrying back-
loads with tump-lines. This means of transportation is still used by
the Atayals of Formosa and the Ainu.
This fatal head-crushing has had an important aftereffect which
the unidentified murderers could not have anticipated. The two
women have gone down in history as a Melanesian (No. 102)
and an Eskimo (No. 103), as stated in dozens of textbooks. This
474
Sinanthropus and the Mongoloids
conclusion is based on a preliminary interpretation of the un-
restored dimensions of the crushed skulls, as a careful reading of
Weidenreich’s original paper will indicate.
His suggestion that the male skull (No. 101) looked like that of
an Ainu was equally hasty. He stated that he made this compari-
son on the basis of some photographs and measurements sent him
by S. Kodanei of Tokyo. At the time S. Kodama’s monumental
work on the craniology of the Ainu 2 had not yet been published.
If we compare the dimensions of No. 101 as given on Table 37
with those of four long series of Ainu skulls from Hokkaido,
Sakhalin, and the Kuriles, we find many differences. The cranial
length of No. 101 is 16 mm. greater than the greatest Ainu mean
and 1 mm. outside all their ranges. The minimum frontal of the
Upper Cave skull is 11 mm. greater than any Ainu mean, and the
bigonial diameter 10 mm. greater, and both these dimensions
fall just inside the maximum ranges of the Kurile and Sakhalin
Ainu, who have the largest faces of all the Ainus. The biorbital
diameter is 9 mm. beyond any Ainu mean, and the nose height
5 mm. beyond any Ainu mean. In both these measurements No.
101 exceeds all Ainu ranges.
The old man of the Upper Cave does not conform strictly to a
Mongoloid model, but neither do all Chinese alive today. In some
respects he resembled the large-faced tribes of American Indians,
like those still living on the Plains. This is particularly visible in
the upper part of the nasal skeleton, and the lateral borders of
the orbits, but the malars and the lower part of the nasal skeleton
are fully Mongoloid in the Eastern Asiatic sense.
The faces of the two female skulls resemble his in general but
are fully Mongoloid in those respects in which his deviates from
the Eastern Asiatic pattern. No. 103, which Weidenreich called
Eskimoid, is the most exaggeratedly Mongoloid of the three. To do
Weidenreich’s memory justice, I will make two brief quotations.
“The Old Man of the Upper Cave appears to represent not only
a very primitive form of modern man but at the same time also a
type of primitive Mongolian.” 3
“. . . the three individuals of the Upper Cave show certain
2 S. Kodama: Crania Ainoica (Sapporo, 1940).
s Weidenreich: “On the Earliest Representatives . . . ,” p. 168.
The Specimen from Kaito-Tung Cave, Teipin, Kwangsi 475
common features in spite of disconformities in others. The former
refers especially to the configuration of the face, namely the low-
ness of its upper part, the quadrangular form of the orbits, the
wide inter-orbital breadth, the shape of the nasal aperture and the
character of its entrance, and the existence of prognathism.” 4
It may be added that although the teeth of No. 101 are not
large, their smallness is largely due to extensive wear, including
interproximal attrition, which reduces the mesio-distal diameters.
The women, who were young when killed, had larger teeth, and
the molars of all three were taurodont. In No. 101, which alone
has a complete dentition, the proportion of molar tooth size to
incisor size is Mongoloid, and Flower s Index, the ratio between
the mesio-distal length of the cheek teeth and the basion-nasion
length, is only 35.7, which is very low, reflecting in part the
extensive wear and in part the exceptionally long basion-nasion
diameter (112 mm). His mandible bore one of Weidenreich’s
criteria linking Sinanthropus with the Mongoloids, a mandibular
torus.
In sum, the Upper Cave skulls from Choukoutien approach
the end of the Sinanthropus-Mongoloid line, bearing the same
kind of relationship to the modern Chinese that the Upper Paleo-
lithic skulls of Europe do to modern Europeans. The sooner we
forget about the Ainu-Melanesian-Eskimo label the better.
The Specimen from Kait o-Tung Cave, Teipin, Kwangsi 5
I n 1956, a fossil human skull base and three tools were found in
a limestone cave in a hill named Chilinshan, in the Leipin District
of Kwangsi. This is the same province from which the enigmatic
remains of Liu-Kiang were recovered. As the fauna was all recent,
there is some doubt whether this specimen is of Late Pleistocene
or post-Pleistocene date. One tool was a crude pebble chopper.
The other two, which were flakes, have not been described.
Found were a combination of palatal and maxillary bones, with
four molars and three premolars; one each of the molar and pre-
4 Ibid., p. 169.
5 Chia and Woo: “Fossil Human Skull Base of Late Paleolithic Stage from
Chilinshan, Leipin District, Kwangsi, VP, Vol. 3 , No. 1 (1959), pp. 37 9.
476 Sinanthropus and the Mongoloids
molar teeth is represented, although all are badly worn, and the
premolars have probably been broken. The palate was medium
to narrow in width; the nasal aperture wide; and the molars within
the size range of the Upper Cave specimens.0 What is left of the
occipital bones is modern in size and form. Otherwise this skull is
too badly broken to indicate much about its racial affinities. Chia
and Woo feel that the form and direction of the stub of a zygo-
matic process remaining on the occipital bone suggest a malar
protrusion of less than Mongoloid proportions, but this seems to
be reading more into the specimen than the evidence warrants.
Post-Pleistocene Skeletons
It would seem that we have carried the northeastern peoples
of the Old World through the Pleistocene in sufficient detail to
show a genetic continuity from Mindel II to the end of Wurm.
The people who live in this area today are Mongoloid, as were the
Chinese from about 3000 b.c. to modern times. The gap of 5,000
years which remains between 8000 b.c. and 3000 b.c. hardly needs
filling, if, indeed, some of those finds most recently described do
not fit into that period. We no longer need to scrutinize each scrap
of bone nor to measure each tooth to a tenth of a millimeter. As far
as northeastern Asia is concerned, our job is done, except for the
problem of the Ainu, who are a white-skinned somewhat Cau-
casoid-looking people with as much hair as a hairy Scot or Jew.
Aside from the recent work of Suzuki, the early man finds in
Japan are limited to a few skeletons from the Jomon Period, a
Mesolithic-Neolithic ceramic culture which has been given an
initial C-14 date of 7500 ± 400 b.c.6 7 I have seen the earliest Jomon
skull in Japan; this skull, like the rest of them, would look better
on the neck of a modem fisherman from Osaka than on that of an
Ainu. However, we do not know what the Ainu were like in
6 Palate width = 37 mm.; nasal opening width = 31 mm.; upper first molar is
10 mm. long and 12 mm. wide; upper second molar = 10 mm. X 13 mm.; upper
third molar = g mm. X 10 mm.
7 H. Befu and C. S. Chard: “Preceramic Cultures in Japan,” AA, Vol. 62, No. 5
(i960), pp. 815-49. (M-769)
America: the Western Extension of the Mongoloid Realm 477
6450 b.c. All the known Ainu skulls are recent. From Manchuria 8
there are a couple of undated, probably Mesolithic or Neolithic,
skulls, which are Mongoloid, and that is all.
America: the Western Extension of the Mongoloid Realm
Both the American Indians and the Eskimo, which inhabit
North and South America, are Mongoloid. All the skulls and bones
of their ancestors which have been unearthed to date are also
Mongoloid. There is not a real Australoid, Melanesian, Negroid,
or Caucasoid piece of bone in the lot.
Three problems concerning the American Indians face us: How
long ago did their ancestors begin to cross the broad glacial plain
of what is now the Bering Strait? Was it initially crossed by people
who could be called H. erectus, or only by H. sapiens? Did the
incursions of Caucasoids, if there really were any, which may
have produced the Ainu and the bearded tribes of the Amur Biver
country and points north, contribute to the peopling of the Ameri-
cas?
First of all, the Bering Strait “highway” to America was not
always open for foot traffic. As Fairbridge’s study of Pleistocene
sea levels showed, it could have been open during the peak of
the Riss-Illinoisan, and again during most if not all of the Wiirm-
Wisconsian (see Chapter 8, p. 314). Theoretically, bands of
hunters living on a Sinanthropus cultural level, with tools good
enough to fashion weapons adequate for killing deer, with fire,
and with a built-in cold adaptation as good as that of the living
Alakalufs (see Chapter 2), could have made the crossing if
they had adequate shelter at night, with or without clothing.
Homo erectus could have done it, but were there any popula-
tions of his grade in the north as late as the height of the Riss-
Illinoisan? The only skull we have in East Asia that can have
come from that period is Mapa, which is so incomplete that we
are not sure whether or not it had crossed the erectus-sapiens
threshold. If people like Mapa crossed at that time, they could
8 A. S. Loukashin: “Some Observations on the Remains of a Pleistocene Fauna
and of the Paleolithic Age in Northern Manchuria,” in G. G. MacCurdy, ed.:
Early Man (Philadelphia: J. B. Lippincott Company; 1937), pp. 327-40.
478
Sinanthropus and the Mongoloids
.
j
i
i
have brought genes for a very archaic skull vault, but probably
also a brain of sapiens size.
No archaeological evidence has yet been unearthed on either
the Asiatic or the American side of the Strait to indicate a Riss-
Illinoisan emigration. The only facts that favor such a migration
are typological. In Venezuela an industry of choppers and
chopping tools has been found in association with extinct animals,
including mastodon, glyptodon (a giant armadillo), mega-
therium (a giant mammal related to the sloths and ant-eaters),
and macrauchenia (a giant three-toed ungulate). Despite the
archaic nature of this fauna, the Carbon- 14 date is only 16,375 —
400 b.c. (No. 0-999), but that is probably the oldest valid date yet
obtained in the New World.9 Junius Bird, who has done con-
siderable excavating near the tip of South America, has found no
evidence of human occupation older than 876 ± 300 b.c. ( W-915 ) .
At that time the Magellanic Indians coexisted with a number of
clumsy old-fashioned mammals like the megatherium, who would
have been extinct in that limited area before that time had anyone
been there to hunt them. If in 8,000 years Indians spread from
Venezuela to the Strait of Magellan, it certainly would not have
taken their ancestors 100,000 years to have gone from Bering
Strait to Venezuela, and 100,000 years ago is the very last date at
which a crossing could have been made over a Riss-Illinoisan
land bridge.
In North America, industries of choppers and chopping tools
have been found in Tennessee and Arkansas underlying mod-
ern American Indian artifacts, but these industries have not yet
been dated. In Maine a similar industry has been tentatively
dated at 2,019 ± 310 b.c.,1 and in the California desert these tools
were made continuously from an unknown date until recent
times.
On these grounds we can probably settle safely for a Wiirm
date of entry, but not necessarily final Wiirm. The Folsom site at
9 I. Rouse: “The Entry of Man into the West Indies,” Y PA, No. 61 (i960),
p. 8; and letter of June 13, 1962. The dating was done by the Humble Oil Com-
pany.
1 D. S. Byers and W. S. Hadlock: “Carbon-14 Dates from Ellsworth Falls in
Maine,” Science, Vol. 121, No. 3151 (1955), pp. 735-6. The date is an average
of two runs, 4150 ± 450 B.P. and 3800 ± 400 B.P. ( M-89).
America: the Western Extension of the Mongoloid Realm 479
Lindenmeier, Colorado, now lias a firm date of 8,820 ± 375 b.c.
(I(UW)-i4i), and the Lehner mammoth site of Arizona, in
which Clovis points were found, one of 9,330 ± 500 b.c. (M-811).
Danger Cave, Utah, a seed-gathering site, is dated at 9,500 ±
600 b.c. (C-609). The Folsom and Clovis industries were ad-
vanced tool-making cultures, certainly not the first in America.
On July 22, i960, The New York Times announced the discov-
ery at Balsequillo, ten miles south of Puebla, Mexico, of a piece of
mastodon pelvis on which someone had engraved sketches of a
bison, tapir, reptile, and apparently a mastodon itself. This carv-
ing had been done when the bone was green. Whether the bone is
really a mastodon pelvis and not a part of some other big animal;
whether or not the drawing really represents a mastodon; and
whether or not it is one of numerous archaeological fakes so com-
monly perpetrated in that country, remain to be determined.
Another lead regarding man’s arrival in America is language.
We observed that in Australia and Tasmania all aborigines speak
or spoke languages of a single family, to which Papuan is probably
also related. On glottochronological grounds, this unity probably
sets a ceiling of 20,000 years on the first settlement of that con-
tinent and those islands. In the two Americas, no one has yet
decided exactly how many linguistic stocks the Indian languages
comprise, but it may well be ten or a dozen. Unless America was
invaded by peoples speaking many languages over a short period
of time, the ceiling of 20,000 years is unnecessary. I believe that
we can postulate with safety that America was first settled some
time in the second half of the Wisconsian (or Wiirm) glaciation,
contemporaneously with the Upper Paleolithic peoples of Europe,
at least in their later stages, with some of the Upper Pleistocene
people of China, and possibly before the time of the Upper Cave
people of Choukoutien.
One further body of evidence is the physical remains of early
American Indians, none of which seem to be older than 10,000
years, if any are that old. Their enumeration and description is
readily available in Wormington’s latest edition of Ancient Man
in North America 2 and need not be repeated here, because it is
2 H. M. Wormington: Ancient Man in North America, Fourth edition (Denver:
Denver Museum of Natural History Popular Series No. 4; 1957).
480 Sinanthropus and the Mongoloids
not necessary to prove that they are both H. sapiens and Mon-
goloid.
However, individuals with archaic cranial vaults turn up
now and then in otherwise normal populations, in both North and
South America, and particularly among some of the Fuegians,
notably the Ona. These vaults have sloping foreheads and are low.
Although they have been referred to as Neanderthaloids in the
literature, both Stewart and Neumann 3 have rightly shown that
these skulls are genetic variants in otherwise fully sapiens, Mon-
goloid populations and do not necessarily mean that whole popu-
lations of low-browed people ever entered America by themselves
and were subsequently absorbed. However, that interpretation,
although unlikely, is not completely ruled out as a faint pos-
sibility for which there is no evidence at present.
From the standpoint of Mongoloid history, the dating of the
arrival of the American Indians is important because the Indians,
by and large, are fully Mongoloid in skin texture and color range,
hair form, hair texture, hair distribution, and degree of sexual
dimorphism. As it is hardly likely that these characteristics of
the soft parts, which distinguish the Mongoloids from all other
subspecies, were acquired independently in Asia and America,
the Asiatic Mongoloids must have acquired them by the time the
ancestors of the American Indians had left Asia for America, in
Upper Pleistocene times.
Conclusion
As late as 1955 it would have been risky to endorse Weid-
enreich’s bold speculation that the Mongoloids of the world are
descended, at least in part, from Sinanthropus or similar popula-
tions of pr e-sapiens Asiatic man, some of which became sapiens
during or shortly after the Riss glacial period. So rapidly are new
discoveries being made in China, and also in Japan, that the risk
is now on the other side. The only serious doubt that remains is
3 T. D. Stewart: “American Neanderthaloids,” QRB, Vol. 32, No. 4 (1957),
pp. 364-9-
G. Neumann: “American Indian Crania with Low Vaults,” HB, Vol. 14, No. 2
(1942), pp. 178-91.
Conclusion
481
this: did Sinanthropus alone and unaided undergo the mutations
in the central nervous system, and probably also the endocrine
system, that transformed him from H. erectus into H. sapiens, or
did someone else who had earlier undergone this process assist
him through mixture? The same problem is involved in the transi-
tion from Solo to Wadjak and the living Australians. We may
never know the answer, but we shall be in a better position to
evaluate what evidence there is after studying the other two
quadrants of the Old World.
11
THE CAUCASOIDS
T he Caucasoid Home
In the northwest quadrant of the Old World we have
more skeletal material to work with than in all the others put to-
gether, but still we are faced with gaps and serious problems. For
example, this is the only section of the world in which no skull of
Homo erectus has been found. The oldest ones whole enough for
diagnosis are already sapiens, but they are not as old as the earli-
est erectus skulls from Java, China, and Africa. Yet they are older
than any other sapiens skulls found elsewhere.
One reason for this unique situation may be that we have not
yet located the earliest Caucasoid homeland. In Europe we have
a succession of remains from the start of the Middle Pleistocene
which are apparently Caucasoid. But it is hardly likely that
Europe was the center of Caucasoid evolution because the suc-
cession that we find is disorderly. The changes in tool industries
are in some cases too abrupt to have been the product of local
technological evolution; yet the tools all emerge from a single set
of traditions. By the same token, successive changes in skulls and
long bones, when we have them, reflect incongruities in what
seems to be a single evolutionary line.
North Africa is also a part of the Caucasoid territory, but it
became so only toward the end of the Pleistocene. Western Asia
is also Caucasoid country. It includes Turkey, all the Arab nations
of Asia, Israel, Iran, Afghanistan, West Pakistan, Kashmir, north-
west India, and parts of Soviet Central Asia west of the Tian-Shan
mountain barrier. Has this broadly delimited area been, like Eu-
The Caucasoids
484
rope, Caucasoid from the beginning; or did it, like North Africa,
serve as the Pleistocene home of another subspecies?
Throughout the Middle Pleistocene the inhabitants of Europe,
western Asia, and Africa were culturally unified in the sense that
all three groups made hand axes, but in the Upper Pleistocene this
unity broke down. The European and western Asian successors
of the hand-ax people continued to follow a single tradition in
tool manufacture, whereas the Africans followed traditions of
their own.
Furthermore, the few skeletons which have been found in Pal-
estine, Lebanon, Iraq, Iran, and Uzbekistan belong to the same
racial line, the Caucasoid, as do those of comparable antiquity in
Europe; but the African skeletons are racially different.
In the parts of western Asia where no ancient skeletons have
been found, including Turkey, Syria, the countries of the Arabian
peninsula, most of Iraq, and all of Afghanistan, West Pakistan,
and northwest India, it may be noted that all the modern inhabit-
ants are Caucasoid except those whose ancestry can be traced to
historic invasions (Huns, Mongols, Turks, etc.) or to the slave
trade (Negroes in Arabia and elsewhere). And, of all these coun-
tries, only southern Arabia, which is part of the Ethiopian faunal
region, contains any trace of a relict population or similar ethnic
enclaves which are not Caucasoid.
In southern Arabia hand axes and cleavers of distinctive Af-
rican style are now being found, and there have long been servile
populations of non-Caucasoid appearance which cannot be en-
tirely explained away as the result of the African slave trade.
Southern Arabia may therefore have been an extension of the Af-
rican-Caucasoid zone of contact as early as the Middle Pleisto-
cene, but the contact between southern Arabia and Africa was
probably broken off somewhat later.
The sum of these three lines of evidence — archaeology, the
study of fossil man, and the study of modern racial distribution —
indicates that western Asia, as defined above, and with one stated
exception, was Caucasoid territory during most if not all of the
Middle and Upper Pleistocene. There we hope to find, if not now,
then eventually as we do more digging, that orderly succession of
culture and race which is so far lacking in Europe.
Contacts Between Subspecies and Caucasoid Evolution 485
Possible Contacts Between Subspecies and
Caucasoid Evolution
In zoogeographical terms, western Asia is a nuclear
region because it stands at the crossroads where Africa, Asia, and
Europe meet and where three faunal regions, the Oriental, Ethi-
opian, and Palearctic, come in contact. With the cooling and
moistening influence of the glacial advances and the warming
and drying of the climate during interglacial periods, Western
Asia has seen the comings and goings of many animal species. The
climatic changes that it has undergone were great enough to be
stimulating, from the evolutionary viewpoint, but not extreme
enough to reduce populations quickly or to cause many extinc-
tions. There was no better place in the Old World for men to
evolve in.
The ancestors of the Caucasoids who, as we suppose, evolved
there could have been in direct peripheral contact with frontier
populations of three of the four other subspecies: the Australoid
in India, the Capoid in North Africa, and possibly the Congoid in
southern Arabia if not also in Africa. The Caucasoids did not have
a common border with the Mongoloids, however, unless they met,
as they do today, at the edges of the plains in Assam and Bengal.
Owing to the former northward extension of Australoids in south-
east Asia it is unlikely that Caucasoids and Mongoloids came into
contact in India any earlier than they did in Central Asia.
It is safest to say that during most of the 500,000 years of man’s
known existence the Mongoloids were in a position to exchange
genes with only one other subspecies, the Australoid. The Con-
goids were in possible contact with two (certainly Capoid and
possibly Caucasoid ) ; the Capoids with two ( Congoid and Cauca-
soid ) ; and the Caucasoids with three, as stated above.
This geographical situation gave the Mongoloids the isolation
necessary to retain their extreme racial peculiarities while evolv-
ing from a lower to a higher grade. At the same time it placed the
Caucasoids in a central position in which they could accept
genes directly and simultaneously from the three other subspe-
cies; process these new genes by exposing them to natural selec-
The Caucasoids
486
tion for climate and culture, in a zoologically central area; and
pass the product back to the peripheral populations separately. In
the same way, to a correspondingly lesser extent, the Mongoloids
could deal with Australoid genes.
Peripheral gene exchanges between the five subspecies in their
formative periods need not have been extensive in order to have
stimulated general evolutionary change, i.e., grade-crossing, in
the populations which received the new genes. Had the ex-
changes been much greater than they were, swampings might
have occurred and some lines might have ceased to exist except
in mixture.
Returning to the Caucasoids, these theoretical exercises suggest
that once we have enough information we can expect to find
continuity in the center of their territory and discontinuity on the
peripheries, such as Europe. If a skull now and then turns up
among them which looks Negroid, Australoid, Capoid, or even
Mongoloid, we should not be surprised because owing to the
spatial position of the Caucasoids, in the middle of the Old World
land masses, they should have been the least “pure” of all human
subspecies.
Continuity and Change in the Caucasoid Quadrant
Before going into details about the evolutionary history of
Caucasoid peoples, let us summarize what we are going to find,
because only with the help of a sweeping survey can this compli-
cated sequence of biological and cultural events be understood.
The material can be divided into four consecutive periods:
(1) from the beginning of the Middle Pleistocene to the end of
the Great or Mindel-Riss Interglacial; (2) the Riss glacial period
and the Last or Riss-Wiirm Interglacial; (3) Early Wiirm, a cold
and wet period lasting into the Gottweig Interstadial; (4) Middle
and Late Wiirm, beginning in the Gottweig Interstadial and end-
ing with the last retreat of the Scandinavian ice around 8,000 b.c.
It would make our task much easier than it is if we had an ade-
quate sample of human remains from each of the four periods in
both Europe and western Asia. We could then test our thesis that
western Asia was the Caucasoid cradle land and Europe a
Continuity and Change in the Caucasoid Quadrant 487
side pocket that received new populations from time to time as
weather permitted. But we cannot do this. Only in Europe is the
sequence of human remains adequate for comparison from period
to period. In western Asia only the third period is well docu-
mented. We are lucky to have this material because it docu-
TABLE 25
PRE-WURM FOSSIL MAN REMAINS FROM
EUROPE AND WESTERN ASIA
Country
Site
Period
Remains
Name
Germany
Mauer (Hei-
delberg)
Earliest Mindel
1 mandible
H. heidelbergensis
Steinheim
Great Interglacial
1 cranium
//. steinheimensis
England
Swanscombe
Great Interglacial
1 calva
H. cf. sapiens
France
Font6chevade
Montmaurin
Monsempron
Last or Riss-Wurm
Interglacial
Last or Riss-Wurm
Interglacial
Last or Riss-Wurm
Interglacial
1 calva, 1 frontal
bone
1 mandible, 4
teeth, 1 vertebra
2 persons; #1 =
cranial frag-
ments & mandi-
ble; § 2 = maxilla
H. sapiens
Italy
Saccopastore
Last or Riss-Wurm
Interglacial
1 skull, 1 calva-
rium
Generally cred-
ited to H. nean-
Germany
Taubach
Ehringsdorf
Last or Riss-Wurm
Interglacial
Last or Riss-Wurm
Interglacial
2 teeth
remains 4 indi-
viduals, skull &
long bones
derthalensis
Czechoslo-
vakia
Ganovce
Last or Riss-Wurm
Interglacial
natural casts of
brain, 1 radius,
1 fibula
Yugoslavia
Krapina
Last or Riss-Wiirm
Interglacial
remains ca. 13
individuals 650
± pieces
Palestine
Mugharet al-
Last or Riss-Wiirm
1 lower rt. mo-
(Israel)
Tabun
Interglacial
lar, 1 piece of fe-
mur
ments the interval which we need most, for the third period is a
time of evolutionary discontinuity in Europe.
In Europe period 1 contains a few precious human remains
which indicate that on that continent man had reached the
threshold of the Homo sapiens grade by at least 250,000 b.c. In pe-
riod 2 no substantial change is evident. As far as we can tell, the
same people continued living there, until at the onset of period 3,
The Caucasoids
488
or a little earlier, a new element was added. That new element
was the famous Neanderthal man, who was more primitive mor-
phologically than his predecessors. Either a new group of people
invaded Europe, absorbing the earlier population, or the earlier
population evolved backward, so to speak, into the Neanderthals.
The Neanderthals continued to live in Europe until the Gottweig
Interstadial, when they disappeared, being followed by the peo-
ple of period 4, the Upper Paleolithic Europeans. Since then Eu-
rope has been continuously inhabited by their descendants and
those of later Caucasoid invaders.
Several facets of this sequence are puzzling. The peoples of
periods 1 and 2 were substantially the same, and their cultures
show an uninterrupted continuity. The culture of the people of
period 3 was derived from that of period 2, although with certain
modifications attributable in part to a change in climate. The cul-
ture of period 4 was new in that it was focused around the pro-
duction of blades, made with the elastic ( horn or antler ) punch,
but it was old in that the types of implements used had been seen
in earlier European tool kits. The burin or graver, for instance, so
typical of the Upper Paleolithic, has been traced back to the
Acheulean hand-ax culture of the Second Interglacial.
In the stone-tool industries of Europe there is a considerable
break between periods 3 and 4, and the racial continuity of
European skulls shows a minor break between periods 2 and 3,
and a major one between 3 and 4. It is possible that the Neander-
thals of period 3 evolved uniquely out of the population of period
2, but the Upper Paleolithic people of period 4 could not have
evolved in Europe out of local Neanderthals.
In western and central Asia there seem to be no sharp cultural
breaks; the succession of physical types was apparently more
gradual. The peoples of period 3 included both Neanderthals of a
less extreme form than those living in Europe, and other people,
in Palestine, who were hardly Neanderthal at all, but transitional
between the Europeans of period 2 and the Upper Paleolithic
people. They apparently invented blade tools, and their imple-
ments foreshadowed those of the Upper Paleolithic. It is likely
that the Upper Paleolithic people and their culture originated
somewhere in western Asia at that time.
The Mauer Mandible, or Heidelberg Jaw
489
The Mauer Mandible, or Heidelberg Jaw 1
So famous is the Heidelberg jaw, more correctly but less
popularly known as the Mauer mandible, that it requires little
description. It was found in 1907 in a sand pit in the village of
Mauer, 6 miles southeast of Heidelberg. It lay 78 feet below the
surface in a soil containing the bones of many animals of a Cro-
merian fauna, but no implements. Among the bones were those
of the spotted hyena, Crocuta crocuta, which did not appear
before an interstadial of Mindel, about 360,000 years ago, ac-
cording to the chronology followed in this book. Mauer is there-
fore as old as Sinanthropus and the Ternefine mandibles from
North Africa, which will be described in the following chapter.
Mauer is a large, massive mandible, chinless and equipped with
blunted gonial angles, but it is not the largest lower jaw yet found.
Both the Sinanthropus male, G-i, and Ternefine 3 are larger in
most dimensions, and even more robust. In fact, Mauer’s index of
robusticity ( see Table 38 ) of 48.8 per cent is lower than the figures
for three of four of the Sinanthropus mandibles, four of five early
North Africa mandibles, and that of Wadjak 2, whereas it is about
the same as that of Pithecanthropus (Sangiran) B. Other Euro-
pean mandibles dated later in the Pleistocene were just as robust
as Mauer, or more so. Individual mandibles from New Caledonia
and the Loyalty Islands of Melanesia, whose living inhabitants
are markedly Australoid, match Mauer in all measurements ex-
cept the width of the ascending ramus,2 and in this dimension
Mauer exceeds all other fossil mandibles of any region or date,
and probably all modern mandibles.
1 0. A. Schotensack: Der Unterkiefer des Homo Heidelhergensis (Leipzig,
1908).
A. Hrdlicka: “The Skeletal Remains of Early Man,” SMC, Vol. 83 (1930),
pp. 90-8.
F. C. Howell: “European and N.W. African Middle Pleistocene Hominids,”
CA, Vol. 1, No. 3 (ig6o), pp. 195-228.
2 R. A. Dart: “Australopithecus prometheus and Telanthropus capensis,” AJPA,
Vol. 13, No. 1 (1955), pp. 67-96.
The conventional measurement of the width of the ascending ramus is a
minimum, in the case of Mauer, 53 mm. Boule and Vallois, in 1952 ( Les Hom-
mes Fossiles ), gave a figure of 60 mm., which is a maximum.
490
The Caucasoids
Fig. 65 Mandibles: Krapina J,
Ehringsdorf, Montmaurin, Hei-
delberg. The progression of Euro-
pean mandibles from Heidelberg
(early Middle Pleistocene) to Mont-
maurin (Late Middle or Early Upper
Pleistocene) to Ehringsdorf and Kra-
pina J (Last Interglacial) runs from
very thick and stout to slightly less
so; from a wide ascending ramus to a
moderate-sized one; and from chin-
lessness to the beginnings of a chin.
Montmaurin, though small, is as stout
as Heidelberg. Ehringsdorf has dis-
proportionately large teeth, and Kra-
pina J has condyles flattened by
arthritis, a disease which sorely
plagued the Neanderthals. (Drawings
of Krapina J after Gorjanovic-
Kramberger, 1906; Ehringsdorf after
Virchow, 1920; Montmaurin after
Vallois, 1955; Heidelberg after a
cast. )
The profile of the symphysis rises steeply in a smooth curve
without any suggestion of a chin. The lower margins of the two
branches of the body, underlying the molars and premolars, ap-
pear swollen in a downward and slightly outward direction, and
then curve upward some 8 mm. to meet in the center line. Inside
the symphyseal region, the bone retreats behind the roots of the
49i
The Mauer Mandible, or Heidelberg Jaw
incisors almost in the form of a shelf, and then dips steeply to the
level of the genial tubercles.
The ascending ramus rises steeply from the body, and the wide
coracoid process is inclined a little forward, as if to accommodate
a forward attachment of the temporals. The general size and form
of the whole ramus suggests a short or medium face length. Judg-
ing from the conformation of the areas of muscle attachment, Hei-
delberg man made extensive use of his temporal and internal (me-
dial) pterygoid muscles, but his masseters were not as strongly
developed or placed as far forward as those of Sinanthropus.
According to Howell ( i960 ) there are three mental foramina
on the left side and two on the right. But the cast shows only one
small foramen on either side. There is no mandibular torus. Mauer
differs from Sinanthropus in that it lacks most of the morphologi-
cal features which characterize the latter, and also in the breadth
of its jaw. Mauer’s bicondylar diameter is only 133 mm. to
148 mm. for Sinanthropus. The Heidelberg skull base, therefore,
was much narrower than Sinanthropus’s, and a narrow base is a
sapiens feature.
All the teeth were in the jaw when discovered, but the whole
left row from the first premolar through the second molar was
broken off in cleaning. Although the teeth are not small, they are
all within the length and breadth ranges of modern man, falling
closest in size to those of Australian aborigines. All the molars are
within the Sinanthropus range, but most of the other teeth fall be-
low it. In other words, the emphasis is on the cheek teeth rather
than on the front teeth, as in the Australoid and Negroid denti-
tions and not as in the Mongoloid and Capoid. Of the molars the
second is the largest, the third next in size, and the first the
smallest.
Howell ( i960 ) has furnished information concerning the molar
cusp patterns. The right first is Y-5, the second and both thirds
are +5. The second and probably the third had a sixth cusp. All
the molars are moderately taurodont, but they lack wrinkling,
cingulums, and dental pearls. The incisors and canines show no
evidence of shoveling. On the whole, the Mauer lower teeth re-
semble those of later Europeans, are not notably different from
those of living Australian aborigines or African Negroes, but dif-
The Caucasoids
492
fer in every pertinent detail from those of Sinanthropus and the
living Mongoloids, and from these of the North African jaws of
equal age.
As a single bone, the Mauer mandible belongs to the expected
grade, considering its antiquity, but because there is no Mauer
cranium we do not know to which species, Homo erectus or Homo
sapiens, Heidelberg man belonged. Both the teeth and the narrow
intercondylar width fit a higher grade than the other features of
the bone itself, and both the jaw and its teeth fail to fit into the
pattern of any of the other four lines of human evolution seen
elsewhere in the world. Mauer therefore stands at the base of a
line of its own.
The Steinheim Cranium3 * * * * 8
In July 1936, a female skull was found in a gravel pit at Stein-
heim an der Murr in Wiirttemberg, 12 miles north of Stuttgart. It
was accompanied by many animal bones, but there were no im-
plements in the gravels. Because it was excavated under labora-
tory conditions by professionals there is no doubt that it belonged
with its fauna. Its date is Great or Mindel-Riss Interglacial,
roughly 250,000 years old, two thirds as old as the Mauer mandi-
ble and possibly 110,000 years younger than Sinanthropus. The
Steinheim woman lived during a warm period.
When discovered, the skull was an almost complete cranium —
the oldest yet found anywhere in the world. The basal part of the
occipital bone had been broken away, as is usual in fossil skulls,
but the front part of the base is present. The front part of the max-
illa below the nasal aperture has been peeled, but not wholly re-
moved, and the front teeth lost. Only the six molars and the right
second premolar remain. Owing to the weight of twenty-three feet
of wet earth covering it, the skull was warped and crushed; the
3F. Berckhemer: “Ein Urmenschenschadel aus dem Diluvialen Schotten von
Steinheim an der Murr,” AAnz, Vol. 10 ( 1933), pp. 318-21.
Berckhemer: “Bemerkungen zu H. Weinert’s Abhandlung ‘Der Urmenschen-
schadel von Steinheim,’ ” VGPA, Vol. 2 ( 1937), pp. 49-58.
H. Weinert: “Der Urmenschenschadel von Steinheim,” ZFMuA, Vol. 35
(1936), pp. 413-518.
Howell: op. cit.
The Steinheim Cranium
493
left side, forward of the earhole, had caved in, and much of the
left side of the face had become detached. The skull has not yet
been restored. In a detailed study Weinert tried to allow for
shrinkage and distortion, but his figures must still be taken as ten-
tative. Some of these are given in Table 37. A few of them, includ-
ing the cranial capacity, have been corrected by Howell ( i960 ) ,
who has handled the original. All that I have had to work with
are photographs and a cast.
The length, breadth, and height dimensions and the cranial
STEINHEIM SWANSCOMBE FONTECHEVADE
Fig. 66 Profiles: Steinheim, Swanscombe, Fontechevade. Steinheim and
Swanscombe are the two oldest specimens of Homo sapiens known. Both come from
the Second or Great Interglacial. Both are designated female. Steinheim is nearly
whole, but badly warped, and it has not been restored. Swanscombe consists of
both parietals and the occipital bone. Fontechevade l consists of a skullcap from
the end of the Middle Pleistocene or Early Upper Pleistocene. The configuration of
the forehead is completely modern, and it apparently had no brow ridges.
Fontechevade 2, not shown here, consists of a small piece of frontal, including the
upper rim of the eye socket. It definitely has no brow ridges. (Drawing A after
Weinert, 1936, and a cast; B after Morant, 1938, and casts; C after Vallois, 1949.)
capacity of 1,150 to 1,175 cc- do not differentiate Steinheim from
the Javanese and Chinese H. erectus skulls, but morphologically
it differs radically from all of them. The occiput is smoothly
rounded, as in modern skulls, and the markings of the neck-mus-
cle attachments are slight and set low. Although low, the forehead
is fairly steep, and the brow ridges stand out like a thin, sharp
visor over the orbits. The skull base is narrow; the mastoids small;
and the side walls of the skull are parallel, as in modern crania,
instead of convergent as in the Eastern H. erecti. The maximum
breadth line is situated at a point 80 per cent of the way up from
the earhole. The highest point on the profile line is located above
the earholes, instead of above the mastoids. Bregma, the point
where the frontal and the two parietal bones meet at the top of the
494
The Caucasoids
skull, is located in front of a vertical line drawn over porion ( the
top of the earhole), as in modern skulls. In Asiatic erectus skulls
bregma lies behind this line. Steinheim’s arc-chord indices, in so
far as they can be reconstructed, are also modern.
In Table 26 the internal dimensions of the Steinheim skull, ten-
tative as they are, are compared with those of four female Asiatic
H. erectus skulls. Steinheim’s internal brain case is shorter than
three, narrower than four, and lower than one, of the four. Never-
theless, its capacity is 100 cc. more than any of them, because of
its shape; it is built like a cube instead of like half of a sphere.
In the sagittal section, lateral view, the frontal lobe shows the
downward bending typical of H. sapiens and what is left of the
TABLE 26
INTERNAL DIMENSIONS AND
CAPACITIES OF STEINHEIM AND
OF FEMALE ERECTUS SKULLS
Ear
Length
Breadth
Height
Capacity
Steinheim
156
121 (?)
103 (?)
1,150-75 cc.
Sinanthropus
11
167
128
102
1015
Solo
1
161
130
103 (?)
1035
Solo
6
153
129
109(?)
1035
Solo
10
159
138
100(?)
1060
occipital part of the base shows the same bending. The hypophys-
eal fossa, or sella turcica, seat of the pituitary gland, seems to be
10 mm. long and 6 mm. deep, as in modern European skulls.4 Al-
though we have no information on the meningeal arteries, in all
known respects Steinheim’s brain cast was sapiens.
The only skull of equal or greater age to which the facial meas-
urements of Steinheim can be compared is Weidenreich’s recon-
struction of the female Sinanthropus, No. 11 (see Table 23).
Steinheim’s minimum frontal is 18 mm. more than S-n’s; its face
breadth is about 16 mm. less; and its biorbital diameter 5 mm. less.
These figures indicate a fundamental difference in the relative de-
velopment of the masticatory apparatus and in the position of the
temporal muscle attachments on the frontal bone. Steinheim’s
face is a little shorter than S-n’s, but still long for a modern Euro-
4 According to Weinert’s drawing, which must be considered with caution.
The Swanscombe Cranial Bones
495
pean woman. Her orbits are smaller than S-n’s, and they differ
from the latter’s in another dimension not included in our table —
Steinheim’s orbits are deep on the lateral or outer side of each cup,
whereas those of Sinanthropus are shallow on the outer side and
deeper on the inner side. The nasal dimensions of the two skulls
are the same, but the nasal bones differ greatly. In Steinheim they
are shaped like an hour glass and pointed at the top, and they
meet at an angle. In Sinanthropus they are parallel-sided and
square at the top, and they meet in a gentle curve.
Below the orbits the zygomatic and maxillary bones of Stein-
heim s face are recessed, as in modern Europeans, rather than
swollen, as in Mongoloids and in Sinanthropus. From the cast and
drawings I have very tentatively calculated the three first indices
of facial flatness, as follows: upper index of facial flatness = ca.
24; simotic index = ca. 55; rhinial index = ca. 40. In all three in-
dices Steinheim is typically Caucasoid, so much so that even if
these figures are 10 per cent or more off, the racial diagnosis must
be the same.
The dimensions of Steinheim’s seven upper teeth are given on
Table 39. These molars and one upper second premolar all fall
within the length and breadth ranges of modern men. Only the
mesiodistal lengths of the first and second upper molars are great
enough to be within the Sinanthropus range. All the teeth are
smaller than the means for living Australians, and the third molar
is even smaller than the mean for living Europeans. The first mo-
lar is the largest; the second is the next larger; and the third is
the smallest. All four first and second molars seem to have four
cusps, and the third is reduced and rounded to such an extent that
the cusps are not easy to distinguish. These teeth are also moder-
ately taurodont. Except for this last feature, nothing notable
distinguishes them from those of a modern European woman.
The Swanscombe Cranial Bones 5
The well-known Swanscombe skull consists of three
separate bones, an occipital and both parietals, found at three dif-
5 The basic report on the occipital and left parietal (the first found) is “Report
on the Swanscombe Skull,” JRAI, Vol. 68 (1938), written by a committee of
The Caucasoids
496
ferent times from 1935 onward, in the 100-foot terrace of the
Thames river gravels, in direct association with a Great or Mindel-
Riss Interglacial fauna and a Middle Acheulean hand-ax and
flake industry. The fluorine content of the skull is the same as that
of the animal bones found with it. Like Steinheim’s, its geological
position is impeccable.
The cranial capacity, variously estimated at from 1,275 to
1,325 cc., is in the range of that of modern European women, and
its breadth and height figures are modern. As one might expect of
ancient bones, Swanscombe’s are thick, ranging from 6.5 to 9 mm.
Morphologically they are essentially modern, with a few archaic
features; for example, the foramen magnum is long and narrow,
and the occipital bone is broad at the base ( biasterionic breadth =
123.5 mm.). On the inside, the brain cast of the occipital lobes
and of the cerebellum are sapiens in configuration,6 and the
channels of the middle meningeal artery are full and complex,
although of a pattern rare in modern peoples.
There has been a great deal of speculation about Swanscombe’s
face, but because Steinheim has a face, and because the threshold
between Homo erectus and Homo sapiens lies in the brain, and
not in the face, it is unnecessary.7 We cannot expect Swanscombe
to have had a face like that of a modern London lady, whose
lineaments have been acquired over many millennia of modern
living, and who would find it difficult if not impossible to survive
authors, particularly W. E. LeG. Clark (“General Features of the Swanscombe
Skull Bones” and “The Endocranial Cast”); and G. M. Morant (“The Form of the
Swanscombe Skull”).
For the right parietal: J. Wymer: “A Further Fragment of the Swanscombe
Skull,” Nature, Vol. 176, No. 4479 ( 1955 ) , pp. 426-7.
For fluorine: K. P. Oakley: “Physical Anthropology in the British Museum,”
in D. F. Boberts and J. S. Weiner: The Scope of Physical Anthropology (New
York: Oxford University Press; 1958), pp. 51-3.
6 Howell (i960, p. 221) says: “The cerebellar fossae are small in comparison
with the cerebral fossae,” which is true if the endocranial cast is compared with
that of a typical modern European, but they are not small when compared with
those of a modern Australian aborigine, or even, for example, a Bronze Age skull
from Tepe Hissar, Iran, which is perfectly Caucasoid.
7 As Howell and others have pointed out, there is a large dimplelike depression
on the forward margin of the occipital bone which may be interpreted as an
extension of the sphenoid sinus. As a large sphenoid sinus may be associated with
a general pneumatization of the face, and hence heavy brow ridges, the con-
clusion is that Swanscombe had heavy brow ridges, but even if she did, my
diagnosis remains unshaken.
European Fossil Men of the Early Upper Pleistocene 497
under the cultural conditions in which both of these ancient fe-
males must have lived. The sapiens grade is broad and inclusive,
covering many subgrades and degrees. If the modern Australian
aborigines are sapiens, these women were sapiens too. And calcu-
lated according to the formula for modern Australian skulls, the
cranial capacity of Steinheim is 1,145 cc., very close to Howell’s
figure. If these women were not sapiens, neither are many of the
living female Australian aborigines and New Caledonians,8 whose
skulls Steinheim and Swanscombe resemble in grade, but not in
line.
European Fossil Men of the Early Upper Pleistocene
Neither Steinheim, Germany, nor Swanscombe, England,
were comfortable places to live in during the next to last, or Riss,
glacial period. The descendants of the two women whose skulls
we have just studied must have moved south as the weather grew
colder. Between the two peaks of the Riss was a mild interstadial
with a climate similar to that of today, and at the end of Riss
came a return to warm conditions with the Riss-Wiirm or Last In-
terstadial. In Europe local populations were undoubtedly most
mobile in the regions of greatest climatic change and least mobile
where the climate was the most nearly constant. If the Europeans
of the Last Interglacial were descended from the Europeans of
8 In 1955 W. E. LeGros Clark accepted these skulls, at least provisionally, as
primitive members of Homo sapiens. In i960 Clark Howell discussed the problem
at length without committing himself. He emphasized the archaic traits that
foreshadowed the Neanderthals in both these skulls. Also in i960, W. W. Howells
postulated the first appearance of Homo sapiens at “almost certainly . . . some
150.000 years ago.” S. L. Washburn, in the same number of the Scientific Ameri-
can, wrote: . . the species Homo sapiens appeared perhaps as recently as
50.000 years ago.” As no one in the Anglo-American world knows more about
Steinheim and Swanscombe than these four experts, who are well aware of the
date of the skulls, the disagreement is obviously a matter of how one defines Homo
sapiens.
Clark: The Fossil Evidence for Human Evolution (Chicago: University of
Chicago Press; 1955), pp. 63-6.
Howell: op. cit.
W. W. Howells: “The Distribution of Man,” SA, Vol. 203, No. 3 (i960),
pp. 113-27.
S. L. Washburn: “Tools and Human Evolution,” SA, Vol. 203, No. 3 (i960),
PP- 63-75-
The Caucasoids
498
the Great Interglacial, therefore, we may expect a certain amount
of discontinuity rather than direct regional continuities; and if
new populations came in from Asia we may expect to find some
evidence of anatomical change.
As shown on Table 25, the Last Interglacial is represented in
Europe by eight sites, and in western Asia by a single site, which
contained only one tooth and a fragment of femur. Ignoring west-
ern Asia for the moment, we find that the eight European sites
have yielded five skulls or sets of skulls, not one as whole as Stein-
heim. We have mandibles from three of these sites; teeth from
five; and body bones from only two. The sites of the skulls are:
Fontechevade, Saccopastore, Ehringsdorf, Ganovce, and Krapina.
The mandible sites are Montmaurin, Monsempron, and Krapina.
The tooth sites are Saccopastore, Montmaurin, Monsempron, Ehr-
ingsdorf, and Krapina; and the postcranial bones come from Ehr-
ingsdorf and Krapina alone.
Except that Montmaurin may possibly belong to the Second
Interglacial, that Fontechevade probably dates from the begin-
ning of the Last Interglacial if not from a Riss Interstadial, and
that Krapina may overlap the beginning of Early Wiirm, we have
no inkling of the chronological order of the eight sites. To describe
the remains from each site one by one would only convey a false
picture of an orderly succession. Instead I shall deal with skulls,
mandibles, teeth, and body bones in that order. In this way Font-
echevade, where only skulls were uncovered, comes first, and
Krapina, which represents the greatest store of body bones, comes
last, and we shall not have to backtrack for comparisons.9
Fontechevade
I n 1947 Mile G. Henri-Martin excavated a cave at Fontechevade
in Charente. In the upper levels she found Mousterian artifacts of
a kind characteristically made by Neanderthal men, but no hu-
man remains. Below this level lay a limy crust, which not only ef-
9 This procedure has another advantage — it avoids anticipation of things to
come. Here there will be talk of pr e-sapiens or pre-Neanderthals. We have already
discussed sapiens , and we shall consider Neanderthal when we get to him.
Fontechevade
499
fectively sealed off what lay below but also indicated a consider-
able time gap between the two layers. Under the crust there was a
Tayacian flake industry with a warm fauna, including the extinct
Merck’s rhinoceros, fallow deer, bear, tortoise, and Cyon, a wild
dog now found mostly in southern Asia. With these animal bones
fragments of two skullcaps, Fontechevade 1 and 2 were also dis-
covered. Under the Tayacian level Mile Henri-Martin found
Clactonian flake tools belonging to an industry older than the
Tayacian, but no human remains. Fluorine tests performed by
Oakley definitely tie Fontechevade 1 and 2 to the Tayacian
fauna.1
Number 2, which is the more nearly complete, consists of a left
parietal bone, the upper half of the right parietal, and the upper
part of the frontal. A few scraps that cannot be articulated with
the rest belong to the lower border of the right parietal and to the
occipital. The left parietal contains a hole with depressed edges,
suggesting death by violence, at an age of forty to fifty years;
shortly after death the bones were charred.
If No. 2 was male, the cranial capacity was probably about
M7° cc-> and if female, about 1,460 cc. The bones are from 7 to
9 mm. thick at various places, as in Swanscombe. The skull is long,
broad, and low, and it verges on brachycrany ( round-skull ) , with
an estimated cranial index of 79. Except for its greater breadth of
about 12 mm. (of which we are not completely certain), Fonte-
chevade 2 resembles Swanscombe closely. The biasterionic
breadth (lower occiput) is great— 126 mm. There is, as in Swans-
combe, a depression at lambda, where the two parietals and the
occipital bone meet; this depression is masked, in profile, by the
smoothly curved contour of the parietals. The junction of the pari-
etal and temporal bones was low by modern standards, and the
lines marking the upper limits of the temporal bones were also
low; the minimum frontal diameter must have been great, close to
120 mm. Enough of the frontal bone is present to indicate that the
1 G- Henri-Martin: “Remarques sur la Stratigraphie de Fontechevade,” VAnth,
Vol. 55, Nos. 3-4 (1951), pp. 242-7.
Oakley and C. R. Hoskins: “Application du Test de la Fluorine aux Cranes de
Fontechevade, LAnth, Vol. 55> Nos. 3—4 ( 1951), pp. 239—42.
H. V. Vallois: “The Fontechevade Fossil Men,” AJPA, Vol. 7, No. 3 (1949)
PP- 339-6o.
500
The Caucasoids
skull lacked massive brow ridges. The question of brow ridges in
the Fontechevade population is solved by an examination of the
other and smaller specimen.
According to Vallois (1949, p. 352), Fontechevade 1 “is repre-
sented only by a piece of the frontal 5.5 cm. high and 4 cm. wide,
but it has great interest in that it comprises the region of the gla-
bella and the left supraorbital ridge, with the internal orbital
process of the same side and a small part of the overlying roof. Its
general appearance and its thickness, being inferior to the skull-
cap already described, shows that it derives from another indi-
vidual; this one was also adult. The essential fact is the absolute
absence of a supraorbital torus: the glabella and the brow ridge
are less developed than in the Upper Paleolithic Europeans, or
even the majority of Europeans of today. They recall, in their
general configuration, skulls of female Europeans; there is no na-
sion depression, and the brow ridge does not extend down to the
upper border of the orbit.”
The fact that the two cave dwellers of Fontechevade had
smooth brows has badly shaken some of my colleagues who be-
lieve in unilinear local evolution, because the French cave dwell-
ers of the following period, Early Wiirm, had heavy brow ridges.
However, if we compare these skulls, and Steinheim and Swans-
combe, with the skulls of modern Australian aborigines, our
problem is solved. In any collection of Australian skulls, or in any
living aboriginal tribe, the range of brow-ridge development is
tremendous. Some have bony visors that rival Solo’s, and others
are as smooth-browed as Fontechevade 1. I cannot believe that
everyone living in France in Fontechevade’s lifetime lacked
brow ridges, any more than all Frenchmen do today. Also, aside
from brow ridges, the Fontechevade skulls are similar to those of
their predecessors of the Great Interglacial.
Saccopastore
The Tayacian flake culture of Fontechevade is believed to
have been derived from the Clactonian, which it overlay in that
Saccopastore ^01
particular cave. From the Tayacian came, presumably, the Mous-
terian, which in its pure form differs from the parent industry in
one principal respect. Whereas the Tayacian tool-makers re-
touched the edges of their flakes with taps or blows from a peb-
ble or stick, the Mousterian artificers pressed off fine flakes with a
piece of bone, producing a finer, straighter edge. This second
technique is called step flaking.
Not all Mousterian tool assemblages, however, follow this sim-
ple formula. In some sites Mousterian flakes are found alongside
hand axes of Acheulian tradition; in others the Mousterian re-
*2?™ Saccopastore i, Krapina, Ehringsdorf. The Europeans of
the Last (Third) Interglacial were a variable lot, differing regionally in skull form.
The Ehringsdorf skull from Germany was the most modem-looking. Krapina in
Tugoslavia yielded at least one brachycephalic skull, and the Saccopastores in Italy
with their low, barrel-shaped vaults, foreshadowed the Neanderthal skull form’
(Drawings of Saccopastore i after Sergi, 1944; Krapina after Gorianovic-
Kramberger, 1906, and Brace, 1957; Ehringsdorf after Weidenreich, 1928, and
Klemschmidt, 1931.)
touching technique was applied to Levallois flakes, which had
been struck off prepared cores, from faceted striking platforms.
Although the Mousterian industry, in one form or another, was
characteristic of the cave sites of the Early Wiirm, it began in the
Riss-Wurm or Last Interglacial, and sites of a relatively simple
and unmixed Mousterian variety from that period are particu-
larly common in Italy.
In a gravel pit at Saccopastore, just outside the walls of Rome,
in material deposited by water and containing such tools, the
skull of a thirty-year-old female was found in 1929 by Sergio
Sergi; and in 193d, that of a male aged about thirty-five was dis-
covered in the same pit by A. C. Rlanc and the Abbe H.
502
The Caucasoids
Breuil.2 No. 1, the female, is nearly complete, although the zygo-
matic arches are missing, along with three teeth, and the brow
ridges were cut off by a shovel at the moment of discovery, thus
revealing an extensive frontal sinus. In No. 2 the skullcap is miss-
ing but the right half of the face, all the palate, the right zygo-
matic arch, and most of the right half of the cranial base are pre-
served, and the inside of the cranial base is in excellent condition.
Neither of these skulls is very large. No. 1 has a cranial capacity
of 1,200 cc., close to the figure for Steinheim. The brain of No. 2,
the male, probably was 100 cc. larger, but it is difficult to tell ow-
ing to the absence of the top of the vault. No. 1 had an extremely
low vault, within the Sinanthropus range and even lower than
Steinheim’s. Nevertheless, Sergi’s careful study of the brain cast
shows that the frontal and temporal lobes were similar to those of
modern men, and that the impressions of the cerebellum in the
basal part of the occipital bone are also completely modern in
depth and form. Furthermore, the sphenoidal angle, which indi-
cates the degree of bending of the brain base between the frontal
and temporal regions, is within the modern range. From the
standpoint of endocranial anatomy, No. 2 was fully sapiens, and
so, presumably, was No. x, for the two are enough alike to have
belonged to the same family.
From the standpoint of external anatomy, however, these skulls
look primitive in some respects and simply strange in others. Seen
from above. No. 1 looks streamlined, like a raindrop falling. The
rear profile, from above, looks rounded and even swollen, whereas
the walls of the brain case converge toward the front and the up-
per jaws appear pinched forward in the form of a muzzle. The
side profile of the brain case shows a sloping forehead and
rounded occiput, which curves uninterruptedly to a weak nuchal
crest located below and in front of the rearmost projection of the
occiput. The brow ridges were apparently not very large.
2 S. Sergi: “Craniometria e Craniografia del Primo Paleantropo di Saccopastore,”
RM, Vol. 20-21 (1944), pp. 1-59.
Sergi: “II Secondo Paleantropo di Saccopastore,” RA, Vol. 36 (1948), pp. 1-95.
A. C. Blanc: “Torre in Pietra, Saccopastore, Monte Circeo — On the Position
of the Mousterian in the Pleistocene Sequence of the Rome Area,” NC, 1958,
pp. 167-74.
Saccopastore 503
Seen from in front and from behind, the brain case looks cylin-
drical, like a barrel lying on its side. The mastoids are small, and
the digastric fossae lateral to them are deep, indicating large di-
gastric muscles (the muscles that open the jaw). The foramen
magnum is located in a position normal for modern European
skulls, and the occipital condyles are so oriented as to indicate a
fully erect posture.
At the region of lambda, this skull has no less than eleven sepa-
Fig. 68 Saccopastore Inca Bones. The Last Interglacial skull of Saccopastore 1
is notable for two things: its almost circular profile when seen from behind or in
front, and its Inca bones. An Inca bone is an extra piece of skull vault, separated
from the others by sutures, and lying in the area of lambda, the meeting place be-
tween the two parietal bones and the occipital bone. Inca bones are so called be-
cause they are common in ancient Andean skulls. They are found principally in
Mongoloid crania, in both the Old and New Worlds. The skull of Saccopastore l, a
Caucasoid Italian of the Last Interglacial, has sixteen Inca bones, possibly a world’s
record; eleven major and five minor. Later on, Inca bones were also characteristic
of Neanderthal skulls. No one knows their function. (Drawing after Sergi, 1944.)
5°4
T he Caucasoids
rate Wormian or Inca bones, more than Weidenreich found in any
Sinanthropus skull.
The face is very long, longer than that of the reconstructed Si-
nanthropus female, but not exceptionally broad; and the bizygo-
matic diameter was probably less than the cranial breadth, as in
most modern skulls. In No. 1, which retains one complete zygo-
matic arch, the bizygomatic diameter was only 96.6 per cent of the
cranial breadth, which is a low figure for a fossil skull.
The nasal skeleton is both long and broad, to match the face;
the orbits are large, and the palate is large and rounded at the
tooth line. The nasal bones extend high into the frontal bone, and
their upper border is curved. Seen from the side, the face is very
prognathous, but only in the upper or nasal segment. Below the
nasal aperture the profile is steep. The kind of prognathism seen
in this skull is the opposite of that of Sinanthropus. Below the or-
bits the surface of the malars and maxillae is flat, rather than in-
dented, as in Steinheim, but the two planes so produced stand at
nearly a right angle to each other, instead of being nearly in line,
as in Sinanthropus and the living Mongoloids.
No. 2 is larger than No. 1, and the contours of the skull are less
rounded, probably because No. 2 was a male. Like No. 1 and like
Fontechevade 2, the male skull probably had a cranial index
in the high seventies. No. 2 also has a less rounded tooth line
than No. 1, for his canines and incisors form nearly a straight
line.
As Sergi noted, these two skulls possess a curious combination
of archaic and modern features. The low vault height, the long
face, and the great prognathism of the upper face are characteris-
tics of Homo erectus, but the morphology of the brain and the
area of neck-muscle attachment are modern. Most anthropologists
do not call these skulls sapiens. However, in terms of the criteria
used in this book, they, like their predecessors of the Second In-
terglacial period, had crossed that threshold, yet carried many
features of an earlier grade with them. As for their racial affinities,
the configuration of the eye-nose triangle, the mask, is Caucasoid
like that of Steinheim, but the vault form of the two Sacco-
pastores is very different from that of the earlier cranium.
The Ehringsdorf Remains
5°5
The Ehringsdorf Remains 8
Between 1914 and 1925 a number of fragmentary human re-
mains were found in two neighboring quarries, Kaempfe’s and
Fischers, at Ehringsdorf, near Weimar, in what is now East Ger-
many. They had been secondarily deposited in a crevice between
two layers of limestone, and scraped and rolled on the way from
their original resting place. This aperture had later been filled with
travertine, a stalagmitic material laid down by water, which then
encased them. With them were animal bones of a temperate fauna
and a quantity of plant materials which indicated a climate like
that of today, such as probably existed during the second half of
the Last Interglacial.
The implements found with the human remains 3 4 were Mous-
terian, but not of a simple or homogenous industry, as at Fonte-
chevade and Saccopastore. They included scrapers of Charen-
tian type, derived from Tayacian; small hand axes of Micoque
type, derived from Acheulian; angular scrapers of a style common
to Syria and the Crimea; fine and coarse drills; and crude burins
or gravers. The diversity of these implements and of their geo-
graphical associations suggests that the people who lived at Ehr-
ingsdorf during late Last Interglacial time may have been the
product of mixture between several related populations.
Aside from mandibles, teeth, and a few long bones that will be
described later, we are concerned with a parietal from Fischer’s
3 H. Virchow: Die Menschlichen Skelettreste aus dem Kampfe’schen Bruch
im Travertin von Ehringsdorf bei Weimar (Jena: G. Fischer Verlag; 1920).
F. Wiegers, F. Weidenreich, and E. Schuster: Der Schiidelfund von Weimar-
Ehringsdorf (Jena: G. Fischer Verlag; 1928).
A. Hrdlicka: op. cit.
O. Kleinschmidt: Der XJrmensch (Leipzig, 1931)— Quoted by Behm-Blancke,
1958.
C. U. A. Kappers: “The Endocranial Casts of the Ehringsdorf and Homo
soloensis Skulls,” JAnt, Vol. 71, No. 1 (1936), pp. 61-76.
G. Behm-Blancke: Umwelt, Kultur, und Morphologie des eem-interglacialen
Menschen von Ehringsdorf bei Weimar,” NC, 1958, pp. 141-50.
4 Behm-Blancke states that there were five different strata, each containing
a separate but related Mousterian industry, and that the human remains were
associated with the second industry from the bottom.
The Caucasoids
506
quarry and a broken brain case from Kaempfe’s. All we know
about the former is that “it is a large, oblique fragment of the left
parietal with large portions missing antero-superiorly and postero-
inferiorly. It apparently proceeds from a juvenile, though hardly
a child s skull, is of moderate thickness (maximum 8.5 mm.) and
shows one important feature, which is a marked and nearly cen-
tral parietal eminence, not dull, posterior, and low down as in the
Neanderthalers, but practically like that in modern man.” 5
The specimen from Kaempfe’s quarry consists of a faceless
brain case. It is believed to have been that of a young adult fe-
male, twenty to thirty years old. During the process of deposition
the individual bones came apart and their edges were scraped
and ground, so that they do not fit together. Weidenreich made a
restoration, filling the cracks with plastilene, and the measure-
ments available in the literature are his. Others who have seen the
skull recently think that the gaps were made too wide. Klein-
schmidt, in 1931, published an alternative reconstruction (of the
cast) which differs from Weidenreich’s principally in that it
makes the vault height (auricular) about 8 mm. lower — 113 mm.
instead of 121 mm. This controversy cannot be settled until a
competent anatomist finds time to restudy the original bones.
In any case, this is a large skull, with a capacity of about
1,45° cc. according to Weidenreich and somewhat less if he was
wrong. But it cannot be very much less. Even if the auricular
height was only 113 mm. it is still higher than Fontechevade 2.
The brow ridges are fairly heavy, like those of Steinheim but
thicker, with a depression over glabella and a fairly steep fore-
head. The general morphology of the brain case is modern, with a
humping in the parietal region which appears in both reconstruc-
tions. The maximum breadth of the skull is high on the parietals,
as in modern skulls. The endocranial cast shows a modern condi-
tion in the frontal lobes. The mastoids are modern in size.
There can be no question that the Ehringsdorf skull, although
archaic in some respects, is sapiens ; and in general morphology
it shows a closer similarity to Steinheim and Swanscombe than to
either Fontechevade or Saccopastore. It seems to be on the main
line of Caucasoid cranial evolution.
5Hrdlicka: op. cit., p. 238.
The Stone Brain from Gdnovce, Czechoslovakia
507
The Stone Brain from Gdnovce, Czechoslovakia
I n 1926 a natural endocranial cast of a human brain was found
in travertine, the same marblelike material present at Ehringsdorf,
in a thick deposit surrounding a thermal spring at Ganovce, near
Poprad, Slovakia, at the foot of the Tatra Mountains. With it were
casts of animal bones, and in 1955 casts were also found of a hu-
man radius and tibia.6 Both fauna and geology indicate the mid-
dle of the second half of the Last Interglacial, contemporary with
Ehringsdorf.
The cast itself is nearly whole, and on the left side some of the
bone remained, including parts of the temporal, parietal, and oc-
cipital. Vlcek’s reconstruction indicates a skull with a capacity of
1,320 cc., long, narrow, and lower than any yet studied except
Saccopastore 1 and possibly Steinheim. The occipital lobes are
bun-shaped, with a flattening at lambda. The greatest breadth of
the brain, and of the skull, lies far to the rear. The section seen
from front and rear looks tubular, as in Saccopastore. The brain
stem is centrally located, indicating, if such a conclusion is justi-
fied, a normal position of the head on the cervical vertebrae.
The brainstem itself is long and narrow in section. The cerebellum
is more or less as in Swanscombe, if not more protrusive below the
occipital lobes. There is no Sylvian crest, and the pituitary fossa
measures about 16 by 15 mm., within the outer part of the mod-
ern range.
This brain belonged to a member of the species Homo sapiens,
at a fairly low subgrade; it shows no evolutionary advance over
Swanscombe and it resembles most closely, of all specimens re-
viewed, Saccopastore 1.
6 Vallois: “Un Homme de Neanderthal en Tchekoslovaquie?” L’Anth., Vol. 55
(i95i), PP- 166-9.
Weinert: “Zwei neue Urmenschenfunde,” ZfMuA, Vol. 43, No. 3 (1952), pp.
265-75.
E. Vlcek : “Neandertalskeno cloveka na Slovensku,” SlAr, Vol. 1 (1953), pp. 5-
132.
Vlcek: “The Fossil Man of Ganovce, Czechoslovakia,” JRAI, Vol. 85 (1955),
pp. 163-71.
Vlcek: “Die Reste des Neanderthalmenschen aus dem Gebiete der Tschecho-
slowakei,” NC , pp. 107-22 and plates.
508
The Caucasoids
The Round-headed, People of Krapina
Between 1895 and 1906, K. Gorjanovic-Kramberger and his
associates excavated some 649 shattered pieces of skull, skeleton,
and teeth in a sandstone rock shelter at Krapina in Croatia, now
Yugoslavia. One reason for the minced state of the specimens was
the fact that the excavators removed dangerous overhanging
rocks with dynamite; another that the skeletons had been broken
up at the time of death. Some of the bones were charred, and it
has been claimed but not proved that some of the deceased were
the victims of cannibalism.
The site contained nine archaeological levels, and all the hu-
man material came from the third level from the bottom. The
fauna of all but the topmost level was a warm one, consistent with
a Last Interglacial date. In the top level were the bones of cave
bear, suggesting the onset of Wiirm I, and those of a marmot, a
cold-weather rodent similar to a woodchuck. This last may or may
not have been intrusive. There seems little doubt that the human
bones from Krapina belonged to the latter part of the Riss-Wiirm
or Last Interglacial.
Over a thousand flint implements were removed from the skele-
ton-bearing level, and most of them still rest in the Natural His-
tory Museum at Zagreb, unstudied. The assembly of tools is
called Mousterian, but Skerlj7 8 states that it includes both Acheu-
lean and pre-Aurignacian elements, and Brodar9 believes that
the commonest implements are broad blades and that there are
7 M. de Terra: “Mitteilungen zum Krapina-Fund unter besonderer Beriicksicht-
igung der Zahne,” SVFZ, Vol. 13 ( 1903), pp. 11-31.
F. Gorjanovic-Kramberger: Der Diluviale Mensch von Krapina in Kroatien
(Wiesbaden, 1906).
B. Skerlj: “Were Neanderthalers the Only Inhabitants of Krapina?” BS, Vol. 4,
No. 2 (1958), p. 44.
F. Ozegovic: “Die Bedeutung der Entdeckung des Diluvialen Menschen von
Krapina in Kroatien,” NC, 1958, pp. 21-6.
C. L. Brace: The Significance of the Krapina Finds (Unpublished paper for
Seminar in Primates and Fossil Man, Harvard University, Cambridge, Mass.,
Nov. 20, 1957).
8 Skerlj, in H. V. Vallois and H. L. Movius, Jr.: Catalogue des Hommes Fossiles
(Algiers, 1952), p. 250.
9 S. Brodar: “Das Paleolithikum in Jugoslawien,” Quartdr, Vol. 1, pp. 140-72
The Round-headed People of Krapina 509
also some microliths in the lot. Even if the flints came from more
than one level, the fact that implements of so many types came
from one cave in Croatia occupied during the latter half of the
Last Interglacial indicates either a gathering of different peoples
or cultural evolution in process, or both. We need more informa-
tion to decide.
The Krapina material includes postcranial bones from practi-
cally every part of the body, and over 270 teeth. Of the skulls, only
five are intact enough even to be identified. They are labeled A
Fig. 69 The Mask of Kbapina. In the Caucasoid realm, very few specimens
older than the Neanderthals of Wiirm I have been found with intact or nearly in-
tact faces. The best preserved is the mask of an adolescent from Krapina. It is
notable for its square orbits, its lack of canine fossa, and its peculiar nasal bones.
The suture between nasals and frontal has an inverted V shape, and the right
nasal bone encroaches on the territory of the left. (A seven-eighth view, after
Ozegovic, 1958.)
through E. Only C and D are adult, and only A, C, and D have
been studied.
Skull A is the cap of a brain case belonging to a child of about
three to five years of age. It consists of the frontal bone, the left
parietal, and part of the right parietal. It is large, with a breadth
of over 150 mm., and was undoubtedly brachycephalic. Its walls
are thin, its forehead steep, and its brow ridges weak. The coronal
and sagittal sutures, and also the metopic (frontal) suture, are
open. For the metopic suture to remain open when the skull had
acquired such size is unusual in living populations and unex-
pected in fossil men. Furthermore, the frontal bone shows tubera
5io
The Caucasoids
frontalia, that is, bosses or projections on either side of the fore-
head, as in a modern child. This is the most modern European
skull we have yet considered in this survey.
Skull C belonged to a young and probably female adult. It con-
sists mostly of a face minus the alveolar region and the lower bor-
der of the nasal opening. Although the top of the skull and most of
the base are missing, parts of the frontal, sphenoid, temporal, and
parietal bones extend around the line of greatest breadth. Gorjan-
ovic-Kramberger was therefore able to make a nearly complete
horizontal outline of the whole skull by mirror reproduction. Also,
the curve of the parietals and what is left of the frontal above
glabella can be projected to give a reasonable estimate of cranial
height, which was not impressive.
This skull is of medium size, at least 1,200 cc. when calculated
by a formula for Australian skulls which allows for the brow
ridges. More important, it is brachycranial (short or round-
skulled), with a cranial index of about 83.7. Fontechevade 2 was
probably 79 or a little more, but this is our earliest adult brachy-
cranial skull from anywhere in the world, and it matches skull A,
the fragmentary child’s skull.
Skull D is a composite of many fragments, including an occipi-
tal bone that Loring Brace found among the unidentified pieces in
the Zagreb Museum in 1959. Its shape is the same as that of skull
C., but it is much larger, with a capacity of at least 1,450 cc. and
the extraordinary breadth of 169 mm., which is exceptional today.
Its cranial index of 85.5 is hyperbrachycranial. Skull E, a frag-
mentary child’s headpiece consisting of parts of a frontal and both
parietals, was also brachycranial. All four skulls which are whole
enough to give an idea of head shape are those of round-headed
individuals, like the majority of living Croats.
The face of these Krapina people is best seen in skull C. The
brow ridges are heavy, but divided over nasion. The orbits are
widely separated, and squarish; the amount of facial flatness seen
in the upper orbital and nasal region is about what one would
find among round-headed central Europeans today. Nasion is
highly placed, and the union of the nasal bones with the frontals
is irregular, forming an inverted V. The upper breadth of the nasal
Mandibles of the Europeans of the Last Interglacial Period 511
bones, 18 mm., is moderate; in a separate pair of nasal bones not
connected to any skull, the breadth is 15 mm. In C as in spare
parts of other skulls the nasal region is Caucasoid in structure. Al-
though neither protruding nor particularly large, the zygomatic
and maxillary bones are full below the orbits, and there is no ca-
nine fossa.
Six pieces of maxilla help complete the picture of the faces of
these people. In all six, age can be determined by the teeth; the
ages range from six to twenty years. The maxilla of the twenty-
year-old individual had an alveolar height of 28 mm., which is
within the modern range. The sixteen-year-old’s maxillary frag-
ment was a little prognathous, and in all of them the palate
seemed broad.
Because of the large brow ridges, the low vault, the usual small
mastoids, and a few other archaic features, one cannot say that
the Krapina skulls are fully modern. But they are fully sapiens
and resemble in an over-all way, particularly in that they are
broad-headed, some of the living European peoples. These skulls
are different from the other skulls we have studied. Within the
Caucasoid framework, the pre-Wiirm population of Europe
showed as much regional variation in cranial vault and upper fa-
cial features as the modern European population does today.
The Mandibles of the Europeans of the
Last Interglacial Period
Except for the Heidelberg jaw, which may be three times as
old as the skulls we have just described, pre-Wiirm mandibles are
limited to specimens from four sites, of Last Interglacial date.
These are Montmaurin (Haute-Garonne) 1 and Monsempron
1R. Baylac et al.: “Decouvertes recents dans les grottes de Montmaurin,
Haute-Garonne,” L’Anth., Vol. 54, No. 3-4, pp. 262-71.
Vallois : “La Mandibule Humaine Pre-Mousterien de Montmaurin,” CRAS,
Vol. 240 (i955)» PP- 1577-99-
L. Pales: “Les Neanderthaliens en France,” NC, 1958, pp. 32-7 and plates.
Some workers have recently stated their belief that the Montmaurin jaw is as old
as Steinheim and Swanscombe. B. Kurten: “The relative ages of the Australo-
pithecines of Transvaal and the Pithecanthropines of Java,” in G. Kurth, ed.:
Evolution und Hominization (Stuttgart: Gustav Fischer Verlag; 1962), pp. 74-80.
512
The Caucasoids
( Lot-et-Garonne ) 2 in France, Ehringsdorf, and Krapina. The di-
mensions of these mandibles are given on Table 38, alongside
those of Heidelberg.
The Montmaurin specimen is a nearly complete mandible
found in 1949 in a cave shaft called La Niche in the complex of
caves at Montmaurin, along with implements identified as Early
Mousterian. It is a small jaw, much smaller than Heidelberg,
which it resembles morphologically, although its bicondylar
breadth exceeds Heidelberg’s slightly, indicating perhaps a some-
what broader skull base. It would be small even for a modern jaw,
and it is completely chinless and very robust. Its lower border is
convex so that, like certain modern Maori mandibles from New
Zealand, it rocks when placed on a table. Vallois says that it has
multiple mental foramina, but in Pales’s picture of the left side,
only one large and one very small foramen are visible.
The Monsempron mandible was one of ten scraps of human re-
mains recovered from a site credited to the Last Interglacial,
a judgment which is not completely certain as some reindeer
bones were found in the site. On the other hand, the artifacts are
called Early Mousterian or pre-Mousterian by Vallois.3 The other
scraps were pieces of vault too fragmentary to merit close study
and a piece of maxilla which was not prognathous and which must
have occluded with its missing lower jaw in a modern-style over-
bite, the upper incisors and canines covering the lower ones when
the jaws are closed. In other fossil specimens the teeth of the two
jaws meet edge-to-edge.
The mandible is a piece of the alveolar edge containing a ca-
nine and two premolars. It had a torus nmndibularis, like the
Sinanthropus mandibles and other Mongoloid jaws of various
periods, and like those of some modern European peoples who live
above the Arctic Circle, particularly in Scandinavia.
The Ehringsdorf collection includes one adult mandible, and
one mandible of a child some ten years old. The adult jaw is
nearly complete. In length and breadth it is virtually identical
2 Vallois: “Les Restes Humaines du Gisement Mousterien de Monsempron,”
APa, Vol. 38 ( 1952), pp. 100-20.
J. Piveteau: Traite de Paleontologie VII (Paris: Masson et Cie.; 1957), pp.
482-3.
3 Vallois and Movius: Catalogue des Hommes Fossiles, Section 88, p. 146.
Mandibles of the Europeans of the Last Interglacial Period 513
with Heidelberg, but it is not as high at the symphysis nor as thick
anywhere. Its principal morphological difference from Heidelberg
lies in the chin region. Instead of a smooth curve, its lateral pro-
file is a double curve, quite sloping in the upper or tooth-bearing
segment, and steep below. It looks as if the teeth were too large
for the bone, and it almost has a chin. In this respect it resembles
Wadjak 2.
The child’s mandible is a right half, with the left side extend-
ing around past the canine. Except for the condyle, the left as-
cending ramus is complete. It is steeper than the adult jaw, with
a 55° angle of inclination compared to the adult’s 38° angle, and
lacks the forward projection of the upper portion. Despite its
tender age, the body is almost as high as the adult’s (28.5 as com-
pared to 31 mm.) It looks more modern than the adult jaw, but
this difference may be due to age, sex, individual variation, or a
combination of these factors.
Krapina has yielded eleven mandibles or scraps thereof, nine
designated by the letters A to J (there is no I) and two that were
illustrated in Gorjanovic-Kramberger’s plates but not designated.
Only F, G, H, and J are adult and complete enough to warrant in-
clusion in Table 38.
All eleven conform to a single general pattern and resemble in
grade and line the two French mandibles and the mandibles from
Ehringsdorf. They go together in the same sense as do the Sinan-
thropus mandibles, and those from Ternefine and other sites in
North Africa. All are thick, prognathous, mostly chinless, and
rounded or blunted in the region of the gonial angle. However,
they vary among themselves, as might be expected in any popula-
tion.
H and J are considerably larger than E, F, and G; the first two
may have been masculine and the other three feminine. In F the
sockets of the four incisiors form a straight line between the ca-
nines, instead of the usual flattish arch. All but two of the mandi-
bles have a single, large mental foramen on each side ( if the area
has been preserved ) ; G has two foramina on the right side and
one on the left, H has two on the left and one on the right. All
the Krapina mandibles are thinner and more slightly built than
Heidelberg, and only J approaches it in size; but J’s coracoid proc-
5i4
The Caucasoids
esses are higher, narrower, and arched backward, whereas Heidel-
berg’s incline slightly forward. Furthermore, all the Krapina man-
dibles are flat in frontal profile, as if in anticipation of a chin.
The Teeth of the Europeans of the Last Interglacial
We have available for study 192 permanent and 28 milk teeth
from the Last Interglacial sites of Europe.4 The permanent teeth
include at least two specimens of each upper and lower tooth.
From Krapina come 144; the other 48 are from Saccopastore,
Montmaurin, Monsempron, Ehringsdorf, and Taubach.
These teeth (see Table 39) constitute a single population in
respect to size. All are within the size range of living peoples, al-
though some are larger than those of modern Europeans. The
teeth of both Heidelberg and Swanscombe could, however, be in-
cluded in this collection. We may therefore conclude that from
the beginning of the Middle Pleistocene to the end of the Last
Interglacial, no substantial change took place in the crown di-
mensions of European teeth, and not much has taken place since.5
However, some changes may have taken place in the relative
sizes of the three lower molars. In Heidelberg the order of size is
different in each side of the jaw. On the right side the second
molar is the largest; the third molar is next largest; and the first
is smallest. On the left side only one tooth, the third, can be meas-
ured, and it is smaller than the first molar of the other side.
Montmaurin has the very primitive order of three-two-one. In
the jaw of the Ehringsdorf child, whose wisdom teeth were uncut
and could not be measured, the second molar was larger than the
first, whereas in the adult female from Ehringsdorf the order is
two-one-three for the right side and one-two-three for the left
side. In the Krapina collection the first molars are clearly the
4 All but one of the teeth, a lower first molar from Taubach, near Weimar,
are from sites already mentioned. For the Taubach tooth see A. Nehring: “t)ber
einen menschlichen Molar aus dem Diluvium von Taubach bei Weimar,” VBGA,
Vol. 27 (1895), pp. 573-7.
5 Some lower median incisors from Krapina seem to be larger than the modem
range, but these were loose teeth and may have actually been laterals, in which
case they are unexceptional.
The Teeth of the Europeans of the Last Interglacial 515
largest, and the second and third molars are of equal size. On the
whole, therefore, the teeth of central Europe are more advanced
in the size-order of the lower molars than are the French ones.
Or, if we compare them in time rather than by geography, the
later are more progressive than the earlier ones.
Our information on the morphology of these teeth is uneven.
The Saccopastore molars seem to be taurodont, judging by the
appearance of a broken tooth in a photograph. The Montmaurin
molars are notable in that all three are longer than they are wide.
Among the Monsempron materials, an upper median incisor is
moderately shoveled and has a lingual tubercle at the base. Its
neighbor, an upper lateral incisor, is also moderately shoveled
and has a smaller basal tubercle. The upper canine has two verti-
cal depressions with a ridge between on the lingual side. Both the
first and second upper premolars have single, moderately tauro-
dont roots, as well as small bulbs of cement on the tips of the
roots. It must be remembered that the jaw in which these teeth
are still embedded has a mandibular torus. In the Ehringsdorf
collection, the permanent teeth of the child’s jaw are taurodont
but the teeth of the adult are not. In the child’s jaw, all three mo-
lars (one of which had not yet erupted) have the primitive Y-5
cusp pattern. The Taubach permanent tooth, a lower first molar, is
small, narrow (width = 85 per cent of length), and five-cusped,
as indeed most modern lower first molars are.
Krapina, of course, supplies the most information. The unworn
upper median incisors are shoveled, like that of Monsempron, but
not to the extent found in Sinanthropus or later Mongoloids.
Theirs is a partial shoveling comparable to that found in certain
modern European teeth, particularly among Finns. None of the
Krapina teeth has the I-beam borders, the wrap-around lingual
edge, the barrel-shaped form, or the double-shoveling found in
Sinanthropus and modern Mongoloids, particularly some Ameri-
can Indians. Two of the Krapina upper median incisors have three
basal tubercles on the lingual side, a feature also found in the
teeth of Sinanthropus, and one upper lateral incisor has two such
basal tubercles. Both an upper and a lower canine have ridges on
both edges and a swollen area in between, as in the Monsempron
canine.
The Caucasoids
516
Taurodontism, present in both Heidelberg and Steinheim,
reaches an extreme development in the Krapina premolars and
molars, particularly in the lower third molar. Some are tubular-
rooted, with open ends. One Krapina tooth has a dental pearl —
a feature found in Sinanthropus, Ainus, and Eskimos, among
others. At least one upper first molar has a Carabelli’s cusp, which
is a European feature. The cusp number of the molars varies be-
tween four and five.
The teeth of these Europeans of the Last Interglacial period re-
semble those of their local predecessors in size and general pro-
portions, but they contain morphological features that relate
them in part to the Sinanthropus-Mongoloid line, unless the com-
mon possession of diagnostic Mongoloid features was a coinci-
dence. Let us not forget that during the Last Interglacial the Cau-
casoids and Mongoloids may have met for the first time.
Postcranial Bones of the Last Interglacial:
the Evidence from Krapina
At several of the European sites considered in this section,
bits and scraps of long bones, ribs, and the like have been found
along with the skulls and teeth, but none of them have been care-
fully described except those from Krapina. From what I have
learned of the others from photographs and brief notes, none de-
viates from the Krapina models. Because the Krapina bones were
measured before Rudolf Martin had standardized osteometry, in
certain instances it is difficult to know how Gorjanovic-Kramber-
ger determined his dimensions. He described, or at least men-
tioned, some 232 bones, as given in Table 27.
In general each bone was, of necessity, treated as a separate en-
tity. When, for instance, he discusses a series of cervical verte-
brae, however, we do not know whether they came from different
necks or were all part of a single neck. The same is true of all
other bones that are grouped, like metacarpals and toes. Some
were male, others female; some adult, others juvenile. This treat-
ment limits the value of the study.
The neck vertebrae fall within the modern European range in
517
Postcranial Bones of the Last Interglacial
TABLE 27
POSTCRANIAL BONES FROM
KRAPINA
Vertebrae
20
Os capitatum
1
Fibula
Ribs
20
Metacarpals
3
Os calcis
Scapulae
12
Phalanges, hand
44
Talus
Clavicles
14
Os coxae
2
Cuboid
Humeri
19
Femora
2
Navicular
Radii
11
Patellae
15
Metatarsals
Ulnae
11
Tibiae
3
Foot phalanges :
measurement and form, but they are a little small. The same is
true of the thoracic and lumbar vertebrae, except that some of the
individuals represented suffered from arthritis of the spinal col-
umn, which makes the vertebrae smaller. Other bones confirm
the evidence of the vertebrae that, as Europeans go, these were
small people.
The ribs are modern, except that although the bones are basic-
ally flat in section, the upper borders are thicker than the lower
borders. The shoulder blades (scapulae) are not only modern
but European in detail.
In the scapular spine, a ridge running from the coracoid
process more or less diagonally toward the vertebral border of
the bone, Vallois has found, among modern men, four major
and several minor types, which involve differences in the at-
tachments of the deltoid and trapezoid muscles. In the European
type this spine is narrow at the junction of its outer and middle
thirds. Then it swells out in the middle third and narrows at the
junction of the inner third. In Negroes it is narrow throughout. In
Melanesians it is thick throughout. In Japanese, and presumably
other Mongoloids, it maintains a more or less constant width, but
it is inclined inward and downward at a steep angle in the outer
and middle thirds and then bends up again in the inner third. The
other three types are less steeply inclined, and form straight lines.
The Krapina scapulae resemble Vallois’s first, or European, type.6
The clavicles or collarbones of the Krapina collection are also Eu-
ropean in size and form, but on the slender side.
6 Vallois: “L’Omoplate Humaine,” BMSA, in five numbers from 1928 to 1946;
see Chapter 7.
51 8
The Caucasoids
The humeri also are rather small and slender, but indistinguish-
able from those of modern Europeans, except that in nine ( we do
not know out of how many, the maximum being nineteen), the
olecranon fossa is perforated; that is, the lower arm could be bent
backward at the elbow, as in gorillas and some contemporary
women. This feature has no demonstrable racial significance but
is interesting because of its high incidence. Among the living, very
primitive Caucasoid Veddas of Ceylon, 50 per cent of the humeri
have it. The bones of the lower arm, the radius and ulna, are
also slender, and the ulna is bowed as if for heavy muscular ef-
fort. The ulna from Ganovce is similarly shaped.
A single os capitatum, a wristbone, is important here because
we also have one for Sinanthropus. Krapina’s is large, like those of
modern Europeans; Sinanthropus’s is small, like those of modern
Mongoloids. The metacarpals and phalanges (finger bones) show
no unusual features from a modern European point of view except
that the terminal or nail-bearing phalanges — the last joint of each
finger and of the thumb — were longer, in relation to the other fin-
ger bones, than is usual in living Europeans. As we do not know
which bones belong together, we cannot tell whether Krapina
man had attained the modern European finger-length formula in
which the index finger is longer than the ring finger.
The two pieces of pelvis, one male and the other female, are
completely modern and do not appear to differ in any perceptible
way from those of modern Europeans. However, these pelves are
fragmentary and do not include the upper branch of the pubic
bone, which is peculiar in the later Neanderthals. The femur,
which articulates into the pelvic bones, is distinctive. The head is
set out unusually far from the shaft, and the angle between neck
and shaft is 120 0 in two bones; this figure stands at the lower
border of the modern range. Most people have a more obtuse an-
gle. All fifteen kneecaps (patellae) are also large for the average
of Europeans, but not larger than those of living individuals. Two
pieces of tibia are rounder in section than they would be in mod-
ern Europeans; this indicates that these people lived out of doors
and squatted while resting. Fourteen pieces of fibula are of mod-
ern design.
The foot bones are well represented, and they too are modern
The “ Neanderthals ” of Europe 519
in form and proportions, except that the last or nail-bearing pha-
langes of the toes are a little long in comparison with the lengths
of the other bones.
Despite the technical difficulties resulting from the scrambled
state and unstandardized measurements of the Krapina skeleton,
we have determined that these people were rather small; that
their bones were not especially heavy; that in certain critical fea-
tures, such as the wristbones and shoulder blades, they were defi-
nitely Caucasoid; and that they had achieved the modern Euro-
pean grade in every respect except perhaps in the articulation of
the femur with the pelvis, and in the length of the last joints of
their fingers and toes, which is very variable even in people alive
today. In so far as we are able to interpret the data published
more than a half century ago, these people were early Caucasoids,
who probably resembled some of the marginal Caucasoids of Asia,
like the Veddas and Dravidians, more than they did the more
sturdily built living central Europeans.
The “N eanderthals” of Europe
I n 1680 a German hymn writer named Joachim Neander died in
Bremen, at the age of thirty. During his short fife he had had
seventy-seven hymns published, and he had been honored by
having a small river valley named after him, the Neanderthal,
near Diisseldorf. His family name, originally Neumann, had been
translated into Greek a century earlier. In 1856, in that very val-
ley, a fossil human skullcap was unearthed and it was called Ne-
anderthal man. Although in 1848 a similar skull had been found
in Forbes’s quarry in the Rock of Gibraltar, its importance was
not recognized until 1864, when it was labeled a member of the
Neanderthal group. Since 1856 Neanderthal has become a com-
mon name in many languages and been given to fossil-man re-
mains in Asia, Africa, and even America. The Solo skulls, that of
Broken Hill, and some low-vaulted American Indian crania have
been so tagged from time to time and by various scientists.
In the last century the fame of Neanderthal man has increased.
He is pictured as a crouching, stooping, squat and brutal creature,
with huge jaws and little or no forehead, and a low grade of in-
520
The Caucasoids
telligence. Flesh reconstructions of his face make him look like an
ape. In this guise he has become the prototype of innumerable
cartoons, in which a slant-browed man, clad in a skin, hits a
woman over the head and drags her unconscious body into a
cave. This, the popular image of Neanderthal man, will probably
be with us for decades to come, because it is picturesque, exciting,
and flattering to ourselves. But it is wrong, and so are most of the
elements in the total Neanderthal concept.
This concept stems from the method of taxonomy by which a
species or subspecies is named for the first or “type” specimen
collected and described. This procedure does not take into ac-
count individual and regional variations. Because the original
Neanderthal specimen consisted of only a skullcap and a few long
bones, there was not much to describe, and in 1911 the honor of
being the type specimen was passed on to a nearly complete
French skeleton, that of La Chapelle aux Saints in the Dordogne.
Nineteenth-century anatomists were struck by the heavy brow
ridges and sloping brow of the original Neanderthal. They had
not yet seen the skulls from Trinil, Choukoutien, Solo, and Broken
Hill, nor did they realize, apparently, how heavy-browed and
low-browed individual Australian aborigines can be. When other
skulls with these features were found in many parts of the world,
the name Neanderthal was applied to all of them, no matter where
or when they lived, what kind of tools they made, or what they
were like in other respects. To dub all skulls with salient brow
ridges and sloping foreheads Neanderthal makes no more sense
than to classify everyone with blood type B as belonging to the
same race.
If the concept of a Neanderthal people is to have any validity,
it must be limited in terms of time, space, and culture. Only in this
way can the Neanderthals have formed a population with a gene
pool of its own. Their time span is Early Wiirm or Wiirm I, from
about 75,000 years ago to the beginning of the Gottweig Intersta-
dial, about 40,000 years ago. Its lebensraum was Europe, western
Asia, and central Asia as far east as the Altai Mountains and south
to the Hindu Kush. Its culture was Mousterian, itself a complex of
earlier tool-making techniques.
521
The “Neanderthals” of Europe
Neither the Neanderthal people nor the Neanderthal tool-mak-
ing techniques could have sprung up out of nothing. We have a
somewhat dim picture of the Europeans of the Last Interglacial;
they could have been the descendants of the Europeans of the
Great Interglacial who had crossed the sapiens threshold but had
not advanced very far beyond it. The Mousterian culture had al-
ready come into being during the latter part of the Last Intergla-
cial, as a derivative of the Acheulian, Clactonian-Tayacian, and
the Levalloisian flake techniques.
The Acheulian hand-ax culture extended beyond Europe into
the Arab countries, southern Iran, and India, and also far into
Africa. The Clactonian-Tayacian flake culture was mostly Euro-
pean and Near Eastern; the Levalloisian was concentrated in
western Asia. In northern Europe we do not know how far these
cultures extended because the ice sheets scraped away all traces,
if there were any. In Russia, which was largely unglaciated, we
find a few hand axes along the northern shore of the Black Sea,
and that is all. The entire central Asian realm, from the Volga to
the Altai, may have been uninhabited before the Last Interglacial.
The oldest implements found there are Mousterian, and Mous-
terian sites and surface deposits have been found on both banks
of the Oxus, and east to Tashkent and the mountains. We do not
yet know whether these sites and deposits date back to the Last
Interglacial or merely to Early Wiirm, but it is more logical to sup-
pose the former than the latter, because the Last Interglacial pe-
riod was warmer than Early Wiirm. In one of these periods there
was probably an extension to the east of the Caucasoid geographi-
cal range.
Did not some of these early Caucasoids penetrate farther, cross
the passes in the mountains, and enter the homelands of Sinan-
thropus and the Mongoloids? We do not know the answer, but
very likely they did. The flints from Ting-tsun (see Chapter 8)
have been given a Last Interglacial date, and they are typologi-
cally similar to the Mousterian flints. Of them Bushnell and
McBurney have said: “This industry, in which only the eye of
faith can distinguish the slightest traces of Chopper-Chopping-
Tool influence, is undeniably of general Middle Paleolithic char-
522
The Caucasoids
I
acter in the Western sense.” 7 Let us grant, for the sake of argu-
ment, that Bushnell and McBurney are right. According to this
interpretation of the flints from the Fen Valley, some Cauca-
soids similar to those we have seen in Europe entered central and
northern China from the West, and mixed with the local popula-
tion, and left their tools behind them when they died. If the Chi-
nese population had not yet crossed the erectus-sapiens barrier,
this injection of genes could have given them the chromosomal
equipment to initiate such a transition. Chinese paleontologists
and archaeologists have found no clearly sapiens skeletons in their
country which are older than the Fen Valley flints.
In return, the invaders could have taken over some Sinanthro-
pus-based genes, particularly those that would give them the ca-
pacity to resist the cold of the oncoming glacial winter. Passing
these genes along to the peripheries of their geographical range in
the west, they could have produced the Sinanthropus-like fea-
tures found in some Last Interglacial specimens, such as the man-
dibular torus, shovel incisors, dental pearls, and a degree of facial
flatness not seen in Steinheim. With the onset of the Wiirm cold,
other such people could have infiltrated Europe from the East and
reinforced in the local gene pool, by natural selection, the physi-
cal features that gave them an advantage for survival in the cold.
Such a reconstruction explains the succession of peoples in Eu-
rope from the Last Interglacial period into the Early Wiirm,
without violence to geography, cultural continuity, or genetic
theory.
This hypothesis faces one serious stumbling block, the skulls
from Saccopastore in Italy. They anticipate the Neanderthals
morphologically; they are associated witli a simple, unmixed
Mousterian culture; and they were found in a country far re-
moved from China.
But Italy lies close to Tunisia, with only Sicily and Malta in be-
tween. We know that the Palearctic fauna, including reindeer,
reached Malta, but not Tunisia. A little seamanship of the kind
that carried the ancestors of the Australian aborigines across
Wallacea might also have served to carry a few North Africans to
7 G. Bushnell and C. McBumey: “New World Origins Seen from the Old
World,” Antiquity, Vol. 33, No. 130 ( 1959), pp. 93-101.
The Numbers and Distribution of the Neanderthals 523
Italy, and the North Africans of the Last Interglacial period re-
sembled the Sinanthropus-derived peoples in a number of ways,
particularly in their teeth.
We have good evidence that North Africans went to Spain dur-
ing Wiirm II. Implements of typical Aterian form ( Aterian is a lo-
cal North African industry) — tanged and barbed pressure-flaked
points — have been found in caves in Almeria and Valencia along
with Solutrean points.8 If some North Africans could have crossed
the western Mediterranean to Europe in Wiirm II, others might
have done so earlier.
North Africa, then, is a second potential source of genetic in-
filtration of Europe which could have initiated the Neanderthal
racial complex, if indeed this complex did not simply arise in Eu-
rope out of local genetic materials by mutation, recombination,
and natural selection.
Enough of theory. Let us examine the bones.
The Numbers and Distribution of the Neanderthals
For present purposes the genuine Neanderthals, or Nean-
derthals in sensu stricto, are represented by the skeletal remains
of people who lived in Europe and parts of western and central
Asia during Wiirm I, and in some places a little later; who dwelt
at times in caves, made tools of a characteristic style known as
Mousterian or Levalloisio-Mousterian; and who bore certain ana-
tomical features in common, notably heavy, undivided brow
ridges, small mastoids, pointed, prognathous faces, and large,
projecting noses. Some also had taurodont teeth.
In the fossil-man social register of Vallois and Movius,9 and in
publications dated after 1952, some eighty-two true Neander-
thals, found in forty-two sites, have been listed (see Table 28).
Their geographical distribution follows a distinct climatic pattern.
For the most part they favored the portions of western and south-
ern Europe now lying south of the present-day January frost line,
8 L. Pericot-Garcia : “A New Site with the Remarkable Parpallo-type Solutrean
Points,” CA, Vol. 2, No. 4 ( 1961 ), p. 387.
9 Vallois and Movius: Catalogue des Hommes Fossiles.
524
The Caucasoids
TABLE 28
NEANDERTHAL AND OTHER REMAINS OF
WURM I OR LATER
Site Country and Description
GERMANY
Neanderthal, near Male, 40-50 years; calva and postcranial bones
Diisseldorf
Neuessing, Kelheim, 1 milk incisor
Bavaria
BELGIUM
Bay-Bonnet, Lifege 1 rt. femur, lower end
Engis, Lifege No. 1, baby skull, fragmentary
La Naulette, Namur Mandible, ulna, metacarpal, all fragmentary
Spy, Namur /No. 1, male or female, 35 years; calotte, fragments of
/maxilla, 14 teeth, postcranial bones
JNo. 2, male, 25 years; fragments of maxilla and mandible,
\ 13 teeth, postcranial bones
No. 3, child; tibia and 2 teeth
FRANCE
Bau de l’Aubesier, 1 milk molar
Monieux, Vaucluse
La Chaise, Vouthon, No. 1, calva and three molars
Charente No. 2, child, 4 years; mandible, 3 teeth, parietal, 1 phalange
La Chapelle aux 1 adult male skeleton
Saints, Corr&ze
Combe-Grenal, Dor- Child, 1 fragment mandible
dogne
La Ferrassie, Dor- 6 individuals: No. 1, adult male skeleton;
dogne No. 2, adult female skeleton, skull crushed;
Nos. 3, 4, and 6, infants; No. 4, fetus
Grotte de l’Hyene, 5 individuals, mostly teeth, fragments of mandible and
Arcy-sur-Cure, Yonne maxilla, fibula, and metatarsal
Grotte du Loup, 1 molar tooth, fragments of a parietal
same
Malarnaud, Mont- Male, 21 years; mandible, 1 molar, 1 verteba
seron, ArRge
Marillac, Charente Adult, fragment mandible with 2 teeth
Le Moustier, Peyzac, Male, 18 years; skeleton, complete
Dordogne
Pech de l’Az<5, Sarlat, Child, 5-6 years old; cranium
Dordogne
Le Petit Puymoyen, 4 individuals: No.l, a half mandible with teeth; No. 2, a
Charente piece of maxilla with teeth; No. 3, a piece of mandible with
teeth; No. 4, two isolated teeth (originals all lost)
La Quina, Gardes-Le- About 12 individuals, principally: No. 1, adult female
Pontaroux, skeleton; No. 2, 8-year-old calvarium; No. 3, 10 + pieces
Charente of skull; No. 4, fragment of a mandible; No. 5, various post-
cranial bones
ii
The Numbers and Distribution of the Neanderthals 525
TABLE 28 ( continued )
Site Country and Description
Regourdou, near Las- Mandible, nearly complete, all teeth, various postcranial
caux, Dordogne bones
Vergisson, near Sol- 3 teeth
utr6, Hte. Sa6nne
BRITISH ISLES
La Cotte de St. Br6- 3 individuals: No. 1, fragmentary child’s skull;
lade, Isle of Jersey No. 2, 13 teeth and a piece of tibia; No. 3, 13 teeth
SPAIN AND GIBRALTAR
Banolas, Gerona, 1 mandible, nearly complete, no teeth
Catalonia
Cova Negra de Bel- 1 right parietal bone
lus, J&tiva, Valencia
Pinar, Granada 1 adolescent skull, fragmentary
Gibraltar 3 individuals: No. 1, Forbes’s quarry, adult female skull,
fragmentary; No. 2, Devil’s Tower, 5-year-old skull, frag-
mentary; No. 3, Genista Cave, 1 molar (lost) (Dating is
unknown for all three)
SWITZERLAND
St. Brais 1 upper incisor
ITALY
Circeo (Rome) 3 individuals: No. 1, adult male cranium;
No. 2, adult mandible; No. 3, adult mandible
CZECHOSLOVAKIA
Sipka, N. Moravia 8-10-year-old child, chin portion of mandible
HUNGARY
Subalyuk, Biikk Mts. 2 individuals: No. 1, adult female mandible, 4 vertebrae,
sacrum, 7 limb bones; No. 2, 6-year-old cranium broken
into over 90 pieces, various vertebrae, ribs, and metatar-
sals— said to be Late Mousterian
RUMANIA
Ohaba-Ponor Cave, 1 first phalange of second right toe
Transylvanian Alps
U.S.S.R.
Kiik-Koba, Crimea 2 individuals: No. 1, teeth, hand, feet, tibia, fibula, patella;
No. 2, a newborn infant
Starosel’e, Crimea lj^-year-old child’s skull
Teshik-Tash, Uzbeki- 8-10-year-old child’s skeleton
stan
526
The Caucasoids
TABLE 28 ( continued )
Site Country and Description
Karain, Adala
TURKEY
1 milk molar
Shanidar, Kurdistan
IRAQ
7 individuals: No. 1, adult male skeleton;
Nos. 2, 3, 4, 5, 6, adult skeletons; 1 infant skeleton
Bisitun, near Ker-
manshah
Tamtama, near
Rezaiyeh
IRAN
1 upper incisor, 1 fragment ulna
1 fragment femur
Mugharet al-Tabun,
Mount Carmel
Mugharet al-Skhul,
Mount Carmel
Mugharet al-Zuttiya,
Galilee
Jebel Qafza,
Nazareth
Shukba, Wadi Natuf
Amud Cave, Lake
Tiberias
PALESTINE
1 adult female skeleton, teeth of 4 individuals
Skeletons of 9 adults and 1 child
1 fragmentary cranium, Galilee man
Skeletons of 5 adults and 1 child
Skeletons of 1 adult and 6 children
1 nearly complete skeleton
Ksar ‘Akil, near
Beirut
LEBANON
1 child’s skeleton, “Egbert”
as well as such contemporary vacation areas as the Crimea and the
coasts of Lebanon and Palestine. Very few remains of these men,
or their implements, have been found in colder places. Most of
central and eastern Europe was apparently too cold for them dur-
ing Wiirm I, although some of them lived on the western slopes of
the Zagros Mountains in Iraq and Iran, and just north of the El-
burz and Hindu Kush Mountains in Iran, Soviet Central Asia, and
Afghanistan.
The western European Neanderthals, living in parts of Ger-
many and in Belgium, France, Spain, and Italy seem to have
formed an essentially isolated population, with little if any gene
1
If
i I
The Western Neanderthals 527
flow elsewhere.1 Possibly a thin line of communication led from
Germany across Czechoslovakia and Hungary to the Black Sea
coast, but this was probably genetically inconsequential.
Owing to the glaciated barriers of the Alps and Pyrenees, the
western Neanderthal domain can be subdivided into three par-
tially isolated regional groups, one living in France, Belgium, and
western Germany, a second in Spain and Portugal, and a third in
Italy. Of these the Italian subregion may have been the oldest.
A. C. Blanc has traced the Mousterian in Italy back to the Biss
glacial period,2 where it seems to have evolved locally. Also the
Saccopastore skulls are the most Neanderthaloid of any of the
Last Interglacial specimens reviewed earlier in this chapter.
Because the skulls from Spain and Italy, few as they are, resem-
ble those from France, Belgium, and Germany in most respects,
we can consider the western Neanderthals as a population and
study them as a unit, with regional variability borne in mind.
The Western Neanderthals
Our sample includes the remains of about fifty-five indi-
viduals, but many of them are too fragmentary for detailed de-
scription. Some consist of items like one femur, one milk molar,
and one crushed baby’s skull. Although they have been found
over the span of a century, some are still in private hands, others
have been lost, and few have been competently described. To
bring them all together, to measure or to remeasure them where
necessary, and to treat them statistically would be a monumental
task beyond the scope of this book. The nearest published ap-
proach to such a treatment is Morant’s work.3
Along with Krapina C and Galilee, which belong elsewhere,
Morant measured the following seven skulls of western Neander-
1 Howell: “Pleistocene Glacial Ecology and the Evolution of ‘Classic Neander-
thal Man, SWJA, Vol. 8, No. 4 ( 1952), pp. 377—410.
2 Blanc: “Torre in Pietra, Saccopastore, Monte Circeo . . . ,” pp. 167-74.
3 Morant: “Studies of Paleolithic Man. II. A Biometric Study of Neanderthaloid
Skulls and of their Relationships to Modem Types,” Biometrika, Vol. 2 (1927)
pp. 310-80.
The Caucasoids
528
thals, and a cast of an eighth: Neanderthal, Spy 1, Spy 2, La
Chapelle aux Saints, Le Moustier (adolescent), La Quina 1, La
Quina 2 (a child, and a cast), and Gibraltar 1. To these I have
added La Ferrassie 1, which is the most complete skull we have,
Circeo 1,4 and the child’s skull of Pech de l’Aze,5 making a total of
eleven. These, along with a small, basic bibliography,6 will form
the basis of the following description.
4 1 measured a cast of La Ferrassie 1 bought from the Musee de l’Homme,
Paris, and of Circeo l in the Philadelphia collection.
5 E. Patte: L’Enfant Neanderthalien du Pech de I’Aze (Paris: Masson et Cie;
1957)-
6 The literature on this subject is extensive, but most of it is secondary. The
following works are either original or comprehensive.
S. Alcobe: “Die Neanderthaler Spaniens,” NC, 1958, pp. 1-62.
R. Bay: “Das Gebiss des Neanderthalers,” NC, 1958, pp. 123-40.
M. Boule: “L’Homme Fossile de la Chapelle aux Saints,” APa, Vols. 6 & 7,
pp. 1911-12.
Boule and Vallois: Les Hommes Fossiles (Paris: Masson et Cie; 1952).
M. Fuste: “Morfologia cerebral de un ejemplar neanderthalense procedente de
la cueva de la Carigiielu, en Pinar (Granada),” TIB S, Vol. 15 (1956).
M. Garcia Sanchez: “Restos humanos del paleolitico medio y superior y del
neo-eneolitico de Pinar (Granada),” TIBS, Vol. 15, No. 2 ( i960), pp. 17-72.
F. C. Howell: “Pleistocene Glacial . . .”
Howell: “The Evolutionary Significance of Variation and Varieties of ‘Nean-
derthal’ Man,” QRB, Vol. 32, No. 4 ( 1957), PP- 330-47-
Howell: “Upper Pleistocene Stratigraphy and Early Man in the Levant,” PAPS,
Vol. 103, No. 1 ( 1959), PP- 1-65-
Hrdlicka: “The Skeletal Remains of Early Man.”
A. Leroi-Gourhan: “Etude des Restes Humains Fossiles Provenants des Grottes
d’Arcy-sur-Cure,” APa, Vol. 44 ( 1958), pp. 1-62.
E. Loth: “Beitriige zur Kenntnis der Weichteilanatomie des Neanderthalers,”
ZfRK, Vol. 7 (1938), pp- 13-35-
Morant: “Studies of Paleolithic Man. . . .”
Pales: “Les Neanderthaliens en France.”
Patte: Les Neanderthaliens (Paris: Masson et Cie; 1955).
Patte: L’Enfant Neanderthalien du Pech de I’Aze.
Patte: “L’Enfant du Pech de l’Aze,” NC, 1958, pp. 265-6.
Piveteau: Traite de PaUontologie, VI I.
Sergi: “La Mandibola Neandertaliana Circeo II,” R A, Vol. 41 (1954),
PP- 305-44- „ ,
Sergi: “La Mandibola Neandertaliana Circeo III, RA, Vol. 42 (1955),
PP- 337-404-
Sergi: “Die Neanderthalischen Paleanthropen in Italien,” NC, 1958, pp. 38-51.
W. L. Straus, Jr., and A. J. E. Cave: “Pathology and Posture of Neanderthal
Man,” QRB, Vol. 32, No. 4 ( 1957), PP- 348-63.
F. Twiesselmann: ‘Les Neanderthaliens decouverts en Belgique,” NC, 1958
PP- 63-71.
The Western Neanderthal Crania
529
The Western Neanderthal Crania
As S e r g 1 ( 1958) has remarked, whereas the Europeans of the
Last Interglacial varied considerably in skull form, the Neander-
thals of Wiirm I are much alike. They are in fact so homogenous
that a strong selective agency must have been pruning off deviant
individuals. Data to document this will be found in Table 37.
All had large brains, with capacities ranging from 1,525 to 1,640
cc. in six male skulls and from 1,300 to 1,425 cc. in three female
Fig. 70 From Neanderthal to Nordic in Wurm I. Profiles of the skulls of La
Chapelle aux Saints (A), Shanidar 1 (B), and Skhul 4 (C). Although these three
men were possibly contemporary, their skulls form an evolutionary sequence from
the low-headed, prognathous La Chapelle aux Saints to the high-headed, orthogna-
thous Skhul 4. The difference is geographical. One interpretation is that La
Chapelle aux Saints lived on the periphery of the Caucasoid racial area during
Wiirm I; Shanidar 1 closer to the center; and Skhul 4 nearest the probable center
of Caucasoid evolution. A second interpretation is that La Chapelle aux Saints
shows the most extreme form of cold adaptation and Skhul 4 the least. A third is
that Skhul 4 was the product of mixture with local Caucasoids who had never be-
come Neanderthaloid. All three interpretations have merit. (Drawings after Boule
and Vallois, 1952; Stewart, 1958; and Keith and McCown, 1939.)
ones. The sex difference of 200 cc. is great, and reminiscent of
Sinanthropus. By and large these are more capacious skulls than
the earlier European ones, and Circeo 1 is notably larger than the
Saccopastores, which anticipated the Neanderthal cranial form in
Italy.
Like that of Saccopastore 1, which was a more primitive skull
in many ways, the Neanderthal crania are globular in the rear;
broad, low, widely curved outward over the earholes; and over-
hanging the area of neck-muscle attachment behind. Their fore-
53°
The Caucasoids
heads are sloping and the arc-chord indices of the frontal and
parietal bones are close to the figures for Homo erectus (see Chap-
ter 8, and Table 36), but not the occipital arc-chord index, be-
cause the Neanderthal occiputs are well rounded.
Fig. 71 Why the Neanderthals Were Not Homo erectus: Occipital Views
of Six Skulls. Seen from the rear, the skulls of Homo erectus (A to D) are
pentagonal in form. This is true of all of them, from the earliest (Pithecanthro-
pus 4) to the youngest (Broken Hill). The skull of Saccopastore 1, the first
European skull to show the Neanderthal form (E), and La Chapelle aux Saints
(F), generally taken as the type specimen of the European Neanderthals, are not
pentagonal but circular, flattened at the bases. A circular occipital profile is a purely
sapiens skull form. This, as well as their high cranial capacities, separate the
Neanderthals from Homo erectus. (Drawings of Saccopastore after Sergi, 1955;
La Chapelle aux Saints after a cast; Broken Hill after Pycraft, 1928; Solo 11,
Pithecanthropus 4, and Sinanthropus 11 after Weidenreich, 1943. )
The Western 'Neanderthal Crania
531
It is commonly stated that Neanderthal man could not have
stood or walked erect because his foramen magnum, the hole in
the base of the skull through which the spinal cord passes into the
cervical vertebrae, was slanted backward. But this anatomical
observation is not true; and even if it were true, the position of the
foramen magnum would not have affected his posture.7
The bases of these skulls are very large, as witness La Chapelle
aux Saints, La Ferrassie 1, and Circeo 1, the most nearly intact of
which is La Ferrassie 1.
In Table 29 an unconventional measurement, inion-prosthion,
expresses the maximum length and a conventional one, bimastoid,
expresses the breadth of the bases of these three Neanderthals.
For comparison, the dimensions of the earliest Upper Paleolithic
TABLE 29
SIMPLE DIMENSIONS OF THE
NEANDERTHAL CRANIAL BASE
Inion-Prosthion Bimastoid
La Chapelle 226 142
La Ferrassie 1 220 (?) 147
Circeo I 222 144
Combe Capelle 191 124
skull, Combe Capelle, are also given. The difference of 30 mm. in
length and 20 mm. in breadth show how much larger-based the
Neanderthal crania were than the cranium of Combe Capelle,
which may have been contemporaneous with the last of the Ne-
anderthals in the Wiirm I— II Interstadial.
The combination of a large base and flattened brain case sug-
gests the possibility of artificial deformation, like the cradling of
some modern peoples of the Balkans and of the mountains of the
Near East. But this is unlikely, for two reasons. No single tech-
nique of cranial deformation, intentional or incidental to some
other practice, is likely to have been employed all over western
7 C. Arambourg: “Sur l’Attitude, en Station Verticale, des Neanderthaliens,”
CRAS, Vol. 240 (1955), pp. 804-6.
Patte: Les Neanderthaliens; and Straus and Cave: “Pathology and Pos-
ture. . . loc. cit.
The misconception arose when Boule faultily reconstructed the cranial base of
La Chapelle aux Saints. In La Ferrassie 1 the foramen magnum slants forward.
532
The Caucasoids
Europe for 40,000 years. And in modern, artificially deformed
skulls, the infant and adolescent specimens are more flattened
than the adult ones. In the Neanderthal infant of Pech de l’Aze,
and in the adolescents of Le Moustier and Pinar, there is less
flattening than in the adult skulls. It was not the hand of man, but
natural forces, that flattened the brows of Neanderthal man.
This flattening is combined with another special feature: a re-
markable forward projection of the face, or beakiness, dependent
on the excessive size of the nose. The Neanderthal nose projects
like a prow, influencing everything around and below it. The oft-
cited fact that the brow ridges, particularly those of La Chapelle
aux Saints, form a continuous torus over the nose is due to the
prominence of the nose. What prognathism these skulls possess is
confined to the nasal region. There is little or none of the sub-
nasal prognathism — projection of the alveolar borders of the
palate — seen in Pithecanthopus and Sinanthropus and in many
modern Australoids and Mongoloids. Although grotesquely so, the
nasal and alveolar region of the Neanderthal skulls is Caucasoid.
The Neanderthal face protrudes because of the nose only. The
jaws, which were carried forward by the nasal skeleton, could
have functioned more efficiently for chewing had they been set
two or three centimeters farther back. Inside the Neanderthal
skulls the nasal passages dip downward below the level of their
openings, thus producing an enlarged nasal chamber. In conse-
quence, the maxillae are long, stretched out on the side, and puffy
on the surface. There is no canine fossa. These peculiarities, which
have led some authors to consider the Neanderthals a species
apart, are a structural unit caused by this nasal domination.
The importance of the nose as the prime architect of the Nean-
derthal face has been generally overlooked because in both La
Chapelle, which is too well known, and in La F errassie 1, which is
still virtually undescribed, the nasal bones were missing at the
time of discovery. Both skulls bear scars suggesting post-mortem
surgery. In both it looks as if the brain had been teased out
through the resulting aperture, anticipating, in a clumsy way, the
handiwork of ancient Egyptian embalmers. Such an operation
may have been substituted for the older and more conventional
one of cutting open the skull base to remove the brain, as was done
The Western Neanderthal Crania 533
in Circeo 1. Because La Chapelle aux Saints and La Ferrassie 1
were both to be buried whole, the brain was removed through
the nose.
Despite this mutilation, the nasal margins of the maxillae in
both skulls are preserved, as is the location of nasion. From these
loci we can see that far from being flat-nosed, as in the widely
copied MacGregor restoration of La Chapelle aux Saints, these
noses could have had straight or even convex profiles, as in the
skull of the eight-year-old child from La Quina, and as in Shani-
dar 1 from Iraq. In both those skulls the nasal bones are intact
and the nasal profile was prominent. In Cicero 1 and Gibraltar 1,
the nasal bones were also intact. These “Mediterranean” Neander-
thals, who lived in a milder climate than that of western France,
were less beaky than La Chapelle aux Saints or La Ferrassie 1,
and their upper jaws, from nasion to prosthion, were shorter.
The western Neanderthals, and particularly the French ones,
must have needed big noses for some reason. The nose serves the
purpose, among others, of warming and moistening the inhaled
air on its way to the lungs.8 In most modem populations living in
cold or dry, or both cold and dry climates the nasal opening is
narrow, but narrowness was impossible for Neanderthal because
of the size of his front teeth. A correlation between nasal-opening
breadth and intercanine breadth was established by Schwalbe
seventy-five years ago.9
Recent military research has shown that in very cold climates
it is not so much the lungs but the brain that is in danger of
chilling by inhaled air. The lungs are a long way from the nose.
In arctic populations necks are generally short, skulls broad and
low, and the distance from nose to lungs less than in many long-
necked tropical peoples. In ordinary human heads and necks the
nasal passages are quite close to the arteries that feed blood to
the brain. In a flat-headed, short-necked individual exposed to
intense cold the proximity of nasal passages to these blood ves-
sels could be critical, for the brain must be kept at a constant
8 A. W. Proetz: Essays on the Applied Physiology of the Nose (St. Louis: An-
nals Publishing Company; 1953 ) , 2nd. ed.
9 G. Schwalbe: Lehrbuch der Anatomie der Sinnesorgane (Erlangen: Besold;
1887).
534
The Caucasoids
temperature. It cannot tolerate variations as can the arms and
legs ( see Chapter 2 ) . The size of the nose in the western Neander-
thals, the expansion of the maxillary sinuses, and the forward po-
sition of the nose in reference to that of the brain case may have
had a survival value under conditions of extreme cold without
adequate headgear or protection for the neck.
As the climate grew colder. Neanderthal men may have in-
creasingly needed a large, projecting nasal “radiator,” particu-
larly as there is no archaeological evidence of cultural improve-
ment that would help mitigate the severity of the climate. Their
adaptation was probably anatomical and physiological, that is,
requiring a large caloric intake, like the adaptation of the Alaka-
lufs, who are exposed to much milder conditions.
In Chapter 2 we saw that the Greenland Eskimo, who live in
the shadow of an ice sheet, keep their faces from freezing in part
by a relatively great blood flow, as indicated by the large bore of
the infraorbital foramina of their malar bones. In the Neander-
thals these foramina are also very large. In the left malar of La
Ferrassie 1, the foramen measures 10 mm. by 16 mm. La Chapelle
has two foramina on each side. The largest, on the left, is 8 mm.
by 7 mm. Modern European foramina, which are single, are
about 3 mm. in diameter. Thus, the Neanderthal foramina were
capable of supplying six or seven times as much blood to the face
as those of modern Europeans, whereas the Neanderthal faces
were no more than twice as large. This evidence strengthens the
concept that the peculiarities of the Neanderthal face were adap-
tive, and not simply archaic survivals.
Despite these adaptive features, the Neanderthal faces are es-
sentially Caucasoid. The brow ridges, which form a bar over na-
sion, sweep in a double arc over the eye sockets and trail out far
to the rear on either side. All indices of facial flatness which can
be calculated (see Chapter 8) indicate that the Neanderthals
were the least flat-faced of ancient mankind. The orbits are large,
as in most early men, and they are round. Although the Neander-
thal faces are absolutely broad, they could not have appeared so
in the flesh, for the zygomatic arches are flat rather than bowed.
The facial breadth was, in fact, smaller than the breadth of the
skull, as in modern Europeans. In all other fossil skulls yet
The Western 'Neanderthal Mandibles
535
studied, the faces are wider than the brain cases. This last pe-
culiarity was largely concerned with a disparity in the relative
development of the various muscles which operated the jaw.
The Western Neanderthal Mandibles
Ten adult or nearly adult mandibles are available for study.
They are La Chapelle aux Saints, La Ferrassie x, Spy 1, La Nau-
lette, Le Moustier, La Quina 9, Regourdou, Arcy 2, Circeo 2,
and Circeo 3. Five others, Spy 2, Malarnaud, Marillac, Petit
Puymoyen, and Banolas are either very fragmentary or un-
described ( see Table 38 ) .
Morphologically they are all more or less alike, but they vary
in size. Arcy 2 is large and massive, as much so as the Heidelberg
jaw, but the others are slenderer. La Ferrassie 1 is less massive
than many modern jaws, being almost paper-thin in the gonial re-
gion. Not one of them has Heidelberg’s pecxdiar conformation at
the chin region; indeed, it is not found in any of the Last Inter-
glacial mandibles. La Ferrassie, La Naulette, Arcy 2, and Circeo 3
have rudimentary chins. The most nearly chinless is La Chapelle
aux Saints, the one usually taken as the type specimen.
At least five of these mandibles have mxdtiple mental foramina.
La Chapelle aux Saints, La Ferrassie 1, and La Naulette have two
each on the right side; La Quina 9 has five. Only Malarnaud has
more than one on the left; it has two on each side. When the
foramina are single, they are very large. Because La Chapelle
and La Ferrassie 1 were both apparently right-handed (as will ap-
pear shortly ) , it seems legitimate to wonder whether there might
not be some connection between handedness and the multiple
foramina phenomenon.
The most unusual feature of these jaws is that the coracoid
process rises from the horizontal ramus well to the rear of the
third molar. In this region the jaw looks stretched out, to match
the protrusion of the upper face, which in turn accommodates
the forward thrust of the nose. In La Ferrassie 1 the distance be-
tween the back edge of the third molar and the front edge of the
mandibular foramen ( a perforation of the table of the inner face
The Caucasoids
536
of the ascending ramus) is 37 mm. on the left side and 38 mm. on
the right. In Combe Capelle, the oldest Upper Paleolithic skele-
ton, the figure is 23 mm. for each side. This measurement could
not be taken on the other mandibles, but in all of them the for-
ward edge of the ascending ramus clears the third molar by about
Fig. 72 Caucasoid Neanderthal
Mandibles: Skhul 4, Tabun 3, La
Ferrassie 1, Circeo 3. The Neander-
thal mandible usually illustrated is
that of La Chapelle aux Saints. It is
omitted here because it is nearly
toothless and altered by senile degen-
eration. Also, it is probably the only
completely chinless Neanderthal
mandible. Circeo 3 is large and stout,
recalling Heidelberg in some respects.
La Ferrassie 1 is long and slender,
almost paper-thin in parts of the
gonial region and ascending ramus.
Tabun 2 is shorter, deeper, and more
prognathous; and its Palestinian suc-
cessor, Skhul 4, is virtually modern.
The extreme forward growth of the
European Neanderthal mandibles is
shown in La Ferrassie 1, in which the
third molar clears the ascending ra-
mus by a full centimer. ( Drawings of
Skhul 4 and Tabun 3 after Keith and
McCown, 1939; La Ferrassie 1 after
Pales, 1958, and a cast; Circeo 3 after
Sergi, 1955-)
The Western Neanderthal Mandibles
537
the same distance. In most other human jaws, ancient or modern,
Caucasoid or otherwise, the rear edge of the third molar and the
front of the coracoid process more or less coincide. In the third
molar the coracoid process overlaps the third molar and part if
not all of the second.
On the inner sides of the Neanderthal jaws the muscular relief
is excessive. Loth, who had only La Chapelle aux Saints and Le
Moustier to study, found that both had powerfully developed at-
tachments for the internal and external pterygoids, that neither
had had very strong temporal muscles, and that the areas of
temporal muscle attachment on the skulls extended far to the rear
but not high on the vault. Loth’s observations apply to the other
specimens now available; indeed, to the group as a whole. With
such an elongated jaw, the temporal muscles were at a mechanical
disadvantage when compared with such types as Heidelberg and
Sinanthropus.
In all the jaws the insertions for the digastric muscles are
wide and deep, matching the digastric fossae on the bases of the
skulls; these fossae are situated between the mastoid and para-
mastoid prominences.
Compared with the older mandibles one notes that the basin-
shaped interior configuration is gone, particularly in La Ferrassie
1. The muscles that control the movements of the tongue are given
as much room to work in as in any ordinary modern jaw.
Finally, the mandibular torus, that ridge of dense bone running
on the tongue side from molars to canine, just below the tooth
line, turns up in Marillac, La Quina 9, and Arcy 2. We have seen
it in Sinanthropus and noted that it occurs among arctic peoples
of both the Mongoloid and the Caucasoid subspecies. Like multi-
ple mental foramina, it is associated with life in cold regions.
Whether or not its presence among the Neanderthals is due to a
genetic association, parallelism, or both is an open question.
In conclusion, the western Neanderthal mandibles may be as-
sessed from three points of view: grade, line, and special adapta-
tion. Arcy 2 is quite primitive, reminiscent in some respects of
Heidelberg, and this jaw is probably the oldest of the lot, dating
from the base of Wiirm I. La Ferrassie 1, which may be among the
latest, is the most nearly modern. The group as a whole falls close
538
The Caucasoids
Fig. 73 Mandibles of Skhul 4, La Ferrassie 1, and Heidelberg, Seen from
Above. A. Skhul 4; B. La Ferrassie 1; C. Heidelberg (after a cast). This sequence
illustrates more clearly than the profiles the progression in Caucasoid mandibles
shown in the preceding figures. The Heidelberg jaw is thick; its ascending rami
moderately flaring; and its symphysial region wide fore and aft, receding, and
braced from inside. The jaw of La Ferrassie 1 is extremely long, and its ascending
rami more nearly parallel. The Skhul 4 jaw is virtually modern. Note the marked
racial difference between these jaws and the flaring Mongoloid and North African
mandibles shown in Fig. 62.
The Teeth of the Western Neanderthals 539
to the mandibles of the Last Interglacial as a whole, but in general
is a little more modern. The grade of these jaws therefore fits
their chronological position. The line is patently Caucasoid, un-
less one considers multiple foramina and the mandibular torus to
be peculiarly Mongoloid characteristics. All the jaws complete
enough to judge show special adaptation, not only in the above-
mentioned features but also in the separation of the tooth row
from the ascending ramus. This last feature is merely a part of
the whole complex of the skull in which the nasal apparatus is
projected forward. The special adaptation of both jaw and cra-
nium are toward the bitter cold of western Europe in which the
Neanderthals lived.
The Teeth of the Western Neanderthals
The custom of burying the dead, which the Neanderthals
seem to have invented, insured the preservation of skulls and
bones, but it did little to increase the number of teeth in any
fossil man collection, because, even if unburied bones decay,
their teeth tend to remain intact. Only 138 western Neanderthal
teeth are available for study. I could find published measurements
of only forty-five of these,1 2 and had to measure casts and photo-
graphs to obtain figures for the others. Thirty-two from a cast of
La Ferrassie 1 are reduced by wear, and the two teeth of La
Chapelle aux Saints (a pair of left second premolars) are stumps.
As the western Neanderthals overworked their teeth, only those
of the very young can validly be compared with other series.
Table 39 presents the figures on these teeth. Their minimum
sizes are mostly dictated by degree of wear; their maxima are
true values. They fall in the same general range as those of Heidel-
berg, the teeth of Third Interglacial Europeans, and modern
people. There has been little change in tooth size in Europe from
Heidelberg to the present, except for a few of the Krapina teeth.
The proportions between molars, premolars, canines, and incisors
1 Spy 1 and 2, La Quina H-5, and La Cotte de St. Brelade, Jersey ( Hrdlicka,
1930); Peche de 1’Aze (Patte, 1957); Arcy ( Leroi-Gourhan, 1958); and Circeo
2 and 3 (Sergi, 1954, 1955)-
540
The Caucasoids
are normal for Caucasoids. There is no common pattern in the
molar rows reflecting consistent size gradation.2
It is an anthropological stereotype that Neanderthal man’s
teeth were taurodont, but this is an unwarranted generalization,
H
Fig. 74 The Upper Incisors of
Neanderthals and Other Early
Caucasoids. A. Ehringsdorf; B. Kra-
pina; C. Le Moustier (Neanderthal);
D and E. Arcy-sur-Cure (Neander-
thal); F and G. Tabun l; H and I.
Skhul 7; J and K. Skhul 5; L and M.
Australian Aborigine. A through E
are upper median incisors, labial view;
D through L are upper median inci-
sors, occlusal view; G through M are
upper lateral incisors, occlusal view.
The upper median incisors of the
Europeans of the Last Interglacial
(A and B) and of the Neanderthal
from Le Moustier ( C ) are only mod-
erately shoveled, in European, not
Mongoloid, fashion. Each has only
slightly raised borders and three teat-
like projections at the base. The Ne-
anderthal incisor (D and E) from
Arcy-sur-Cure has a ball-like basal
projection. The median and lateral
incisors from Palestine, those of Tabun
1 ( F and G ) and Skhul 7 ( H and I ) ,
although considerably worn, still show
the effect of moderate shoveling of
the European type. But the less worn
median and lateral incisors of Skhul 5
(J and K) show no sign of shoveling
and closely resemble those of an
Australian aborigine ( L and M ) . This
resemblace corresponds to the Austra-
loid character of Skhul 5’s face.
( Drawings A, B, C after Weidenreich,
1937; D and E after Leroi Gourhan,
1958; F through M after Keith and
McCown, 1939.)
2 The pattern 1-2-3, in which the first molar is largest, the second molar is
next largest, and the third smallest, occurs once; 2-1-3 occurs twice; 2-3-1 four
times; 3-1-2 twice; and 3-2-1 three times.
The Teeth of the Western Neanderthals 541
true of less than half the molars and premolars. And not one of
these is as taurodont as those from Krapina. La Ferrassie 1, La
Quina, Regourdou, and Arcy had no taurodontism at all.
The crowns are also variable. The Dryopithecus pattern, Y-5, is
the commonest, but sixth cusps occur, and the plus-4 pattern is
frequent in upper molars. A cingulum and wrinkling are seen in
one molar from Arcy, and another cingulum is present at the base
of the only unworn Le Moustier canine. Upper incisors from Le
Moustier, St. Brais, and Arcy are moderately shoveled in that they
have raised lateral ridges. A lateral incisor from Le Moustier has
short, fingerlike mesial ridges on the tongue side; the St. Brais
and Arcy specimens have, instead, well-developed basal emi-
nences that make them very thick at the gum level. The Le
Moustier canine with the cingulum also has a spatulate cutting
edge, like that of an incisor, whereas one of the Arcy canines has
a median vertical ridge.
All these special features are reminiscent of Sinanthropus and
the modern Mongoloids in an attenuated way. The cingulum and
wrinkling are also generally archaic, whereas the moderate shovel-
ing is frequent among northern Europeans. We seem to have in
the western Neanderthal series a watering down of the partially
Mongoloid, or pseudo-Mongoloid, dental features of Krapina.
The condition of these teeth gives some idea of the cultural
activities of these cold-weather people. All but the teeth of the
very young are heavily worn, and some are worn in such a fashion
that activities other than chewing food must have been responsi-
ble. In La Ferrassie 1 the outer surfaces of the upper incisors and
left upper canine are polished down, as if by some object like the
corner of a skin or a thong held in the teeth, and in the mandible
there is a gap between the left second incisor and the canine.
These two teeth were twisted away from each other and there is a
pit, as of an abscess, in the outer surface of the mandible, exposing
the root of the left lower canine. There are also similar gaps be-
tween the first and second molar on either side of the mandible.
These dental peculiarities, when added to the evidence for a
special development of the pterygoid and digastric muscles,
suggest that the western Neanderthals softened skins with their
teeth. The abundance of flint tools identifiable as fleshers sup-
542
The Caucasoids
ports this inference. The bitter cold of Wiirm I must have placed
a premium on warm clothing, particularly on serviceable footgear,
as a necessity for survival.
Having good teeth was important to the western Neanderthals;
yet a man did not necessarily die when his teeth were gone. La
Ferrassie 1 had such severe arthritic erosion of the condyles of
his jaw that he could not have chewed his food. La Chapelle
aux Saints had only two teeth; he also was arthritic, and for
many years before his death could not possibly have hunted, nor
could he have chewed roasted meat. Someone must have brought
him his food, and softened it for him. Despite his economic use-
lessness, he was important enough to warrant being buried in a
cave. This could not have happened to everyone who died in a
cave in winter, or the caves would be full of skeletons. During the
lifetime of La Chapelle aux Saints the French Neanderthals were
not poor providers on the verge of death from exposure or starva-
tion. They were competent hunters with some kind of division of
labor based on age and with solicitude for the old and incapaci-
tated.
The Fostcranial Skeletons of the Western Neanderthals
Except for an odd bone here and there, all we know about
western Neanderthal anatomy from the neck down is derived
from seven skeletons: Neanderthal, Spy 1, Spy 2, La Chapelle aux
Saints, La Ferrassie 1, La Ferrassie 2, and La Quina H-5. The
first five are masculine, the last two feminine. Not one is com-
plete. The most fully described is that of La Chapelle aux Saints,
who suffered from disease when alive and whose bones were, as
we shall see, inaccurately reconstructed after exhumation.
Earlier European comparative material consists almost entirely
of the bones from Krapina, which, although more numerous,
were scattered and broken because they had not been buried.
In general, the Neanderthal bones resemble those from Krapina
morphologically but are heavier, as the western Neanderthals
were powerfully built people.
The only vertebral column thoroughly studied is that of La
The Postcranial Skeletons of the Western Neanderthals 543
Chapelle aux Saints, which had been shrunken and distorted by
arthritis and senility. The neck vertebrae are short-bodied, and
he undoubtedly had a short neck, even in his prime.3 So did the
female, La Quina H-5. The thoracic and lumbar vertebrae of La
Chapelle aux Saints are also small, but according to Hrdlicka
( 1930 ) the lumbar vertebrae have unusually large articular facets
on their transverse processes and large neural canals. In both La
Chapelle aux Saints and Spy 2, the top third or fourth of the
sacrum is preserved. In both, the upper margin is narrower than
the modern European mean, but within its range. At least in La
Chapelle aux Saints, the wings of the sacrum rise to a higher level
than the central body. This condition is super-Caucasoid; it is
found in 21.5 per cent of living Europeans and is rare or absent in
other races.
The Neanderthal ribs are known from five fragments from La
Chapelle aux Saints, and the nearly complete rib cages of both La
Ferrassies. These ribs are variable in section. Those of La Cha-
pelle aux Saints were round or triangular, like Krapina’s, but those
of both La Ferrassies were ribbon-shaped, as in modern Euro-
peans. As in Krapina, these ribs all curve so as to produce a deep
chest.
The clavicles of five specimens, Neanderthal, Spy 1, La Cha-
pelle aux Saints, La Ferrassie 1, and La Quina H-5, are longer,
slenderer, and straighter than those of most living Europeans, and
their shape indicates a deep chest. In four fragmentary scapulas
(from the skeletons cited above, excepting La Quina H-5) the
same configuration is seen as in Krapina; but in the Neanderthal
fragment and in La Ferrassie 1 the glenoid cavity, in which the
head of the humerus rotates, is inclined a little more to the rear
than usual in modern specimens, and the ridge for the insertion
of the teres minor muscle is strongly developed. This is the muscle
that rotates the humerus sidewise when the upper arm is held
close to the body. These features are absent in Spy 2, a female,
3 Boule ( 1911-13) made much of the fact that the dorsal spines of this speci-
men’s cervical vertebrae pointed backward instead of downward and backward,
and that the dorsal spine of the sixth cervical vertebra was not bifurcated. As
Arambourg (1955), Patte (1955), and Straus and Cave (1957) have shown,
these features are common in living Europeans.
544
The Caucasoids
who probably did not do the same kinds of work with her arms
that men did.
In six skeletons (all but La Quina H-5) pieces of both right
and left humeri are present, but only the right humerus of Nean-
derthal is complete. His left humerus had been injured early in
life and was underdeveloped. Judging by the relative develop-
ment of his two humeri, La Chapelle aux Saints was strongly right-
handed. Neanderthal’s right humerus is 31.4 cm. long, exactly the
modern European mean. It is a stout bone, but no stouter than
those of many living Europeans.4
In all the western Neanderthal humeri the olecranon fossa, a pit
on the dorsal side of the lower end of the bone which receives the
olecranon process of the ulna when the arm is extended, is large
and deep. It is also perforated in La Quina H-5, both La Ferras-
sies, and Spy 2, making a ratio of perforation of 44 per cent in
nine bones. Virtually the same ratio occurs at Krapina.
We have radii ( the paired lower arm bone on the outer or thumb
side of the wrist ) for all but La Ferrassie 2. Like the humeri, they
are as long as the European mean (22.7 cm. for both sexes). They
are strongly built, and characteristically curved outward, more so
than in any known modern population. Such a curvature pro-
vides a great deal of room between radius and ulna (the com-
panion lower arm bone ) for the development of powerful forearm
muscles. The western Neanderthals must have had, and probably
needed, very strong hands.
The ulnae of these skeletons match the radii in length and stout-
ness, but vary in curvature. That of La Chapelle aux Saints is
nearly straight; La Ferrassie l’s is slightly curved and La Ferrassie
2 s is very curved. In all of them the olecranon process, which is
the peak of the elbow (the “funny bone”) and fits into the ole-
cranon fossa of the humerus when the arm is extended, is excep-
tionally long. This gives the triceps muscle a great mechanical ad-
vantage in extending the forearm.
Nearly complete hand skeletons were found with both La Fer-
4 Boule calculated an index of robusticity by dividing the minimum shaft cir-
cumference by the maximum length of the humerus. The result was a figure of
23 per cent for Neanderthal. The mean for modem white American males is
21.5 per cent, and the upper range far exceeds 23 per cent.
The Poster anial Skeletons of the Western Neanderthals 545
rassie specimens.5 Spy 1 has two metacarpals, Spy 2 six each of
metacarpals and phalanges; and La Chapelle aux Saints has one
fragmentary scaphoid ( the wrist bone articulating with the
thumb side of the radius), one capitate (the central wrist bone
of the outer row articulating with the third and fourth meta-
carpals ) , three metacarpals, and two phalanges, all from his little
used, probably defective, left hand.
Only the two wrist bones of La Chapelle aux Saints have been
described. The scaphoid is said to be small and flattened, but
this description does not apply to the half of a scaphoid present
in La Ferrassie 2’s right hand.6 The capitate of La Chapelle aux
Saints is as long as Krapina’s (24 mm.) but narrower ( 14 mm. as
compared to 18 mm.); La Ferrassie 2’s capitate is of normal size
and proportions for a European woman (22 mm. by 16 mm.), as
are the rest of her surviving wrist bones.7 8
La Chapelle aux Saints’s left first metacarpal (that of the
thumb) is a little short by European standards — much has been
made of this point s — but the corresponding bone of La Ferrassie 2
is longer than the mean for European women. Her metacarpals and
phalanges also fall close to modern female European means, ex-
cept that the terminal phalanx of her right little finger is short
and conical, apparently a congenital defect. A single fourth left
metacarpal from Arcy is also perfectly modern.9 On the whole,
no evidence yet produced indicates that the western Neanderthal
hands were notably different from those of hard-working modern
Europeans.
Scanty but generally adequate specimens are available for most
of the bones of the lower extremity. Among the least satisfactory
are those of the os coxae, or pelvic bone. These consist of one
piece of left ilium from Neanderthal, and two similar pieces, one
5 F. Sarasin: Die Variationen im Bau des Handskeletts verschiedener Mens-
chenformen,” ZfMuA, Vol. 30 ( 1931), pp. 252-316.
6 Piveteau: Traite de Paleontologie, Vol. VII, photograph on p. 458.
7 The lunate and triquetral are missing.
8 Boule also stressed the fact that the proximal articular surface of this bone is
convex rather than saddle-shaped, as in modem metacarpals; but in this respect
both La Ferrassies are normal.
9 Leroi-Gourhan : op. cit., p. 54.
Length is 55 mm. ( ? ) ; knuckle breadth = 14 mm.; minimum shaft diameter =
6 mm. The last measurement is actually small.
The Caucasoids
546
left and one right, from La Chapelle aux Saints. All observers
since Boule 1 agree that they are similar to those of modern men.
Femora are available for Neanderthal, Spy 2, La Chapelle aux
Saints, both La Ferrassies, and La Quina H-5. They are of medium
length ( 44 cm. for four males ) , about as long as those of modern
Bavarians. They are also relatively thick and heavy, like those of
the most solidly built modern populations, for example, the Japa-
nese. The diameter of the femoral head (54.7 mm. for four
males) is great compared to the same diameter in most people,
but not compared to modern Europeans, whose femoral heads are
large. But the western Neanderthal femora differ from the modern
European norm in three respects. They have weakly developed
pilasters on the backs of their shafts; these shafts are strongly
bowed, like those of modern peoples who squat on their heels;
and the angle between the neck of the femur and its shaft is low,
about 1180 for males. Most modern femora of all races have
mean angles of from 121 to 133 °. However, the femoral angle of
the female La Ferrassie 2 was 123 °, or quite modern. What this
angle signifies is not clear.
We also have four tibias, one each for Spy 2, La Chapelle aux
Saints, and the two La Ferrassies. Only that of Spy 2 is complete.
It is 33.7 cm. long, or 78.7 per cent of the length of its femur. This
tibiofemoral ratio is among the lowest known, the same as that
for modern Lapps. Relatively short tibias are characteristic of
circumpolar peoples. Like the femora, the western Neanderthal
tibias are bowed. Their heads are bent back at angles of as much
as 200, like the heads of other notable squatters, the Fuegians
and the California Indians.
Also available are patellas, or kneecaps, for Spy 2, La Chapelle
aux Saints, and one of the odd La Quina skeletons. They are all
perfectly human, and large. Fibulae were found with Spy 1 and
both La Ferrassies, but none has been described.
The feet of the western Neanderthals were almost as distinctive
as their skulls, as we can see from a set of footprints found in an
Italian cave by A. C. Blanc (see Fig. 75). They look somewhat
1 Straus and Cave: op. cit.
Piveteau: op. cit.
TTie Poster anial Skeletons of the Western Neanderthals 547
like the prints of a modern Alakaluf Indian foot used to walking
in icy water and snow, and very unlike the slender, tapering
marks left by an Upper Paleolithic man in another cave in Italy.
The Neanderthal prints are very wide, in heel, ball, and toe, and
the toes are very short, except for the great toe. Despite the ex-
treme proportions, there is nothing about these prints that is not,
as some have stated, completely human.
The available fossil foot bones tell the same story.2 We have a
right calcaneum and a left astragulus for Spy 2; a calcaneum, an
astragulus, and five metatarsals — all seven bones broken and in-
complete for La Chapelle aux Saints; and three nearly complete,
articulated feet for the two La Ferrassies, one of which, the right
foot of La Ferrassie 2, is depicted in a scale photograph,3 which I
measured.
The calcanea are large and thick, with long heel portions, as in
European (and not, as supposed, in Negro) heels. On the under-
side of every calcaneum is a facet called the sustentaculum tali ,
the outermost of the three which hold up the astragulus, or ankle
bone. In the western Neanderthal calcanea this facet is unusually
large, indicating that a great deal of weight fell on the outer half
of the foot. The astragali are short in proportion to height, and the
other metatarsal bones, as seen in the foot of La Ferrassie 2, are
wide and square. La Ferrassie 2’s metatarsals are about 2 mm.
longer than those of European women of today. Also the meta-
tarsals of the Neanderthal woman differ less in length than those
of modern European women; in La Ferrassie 2 the second meta-
tarsal is only 6 mm. longer than the fifth, whereas in European
women the difference averages 10 mm. The proximal phalanges
(the toe joints nearest the body of the foot) are a little shorter
than the modern means, except for that of the great toe, which is
just as long. La Ferrassie 2, therefore, had a foot of normal length
for a European woman, but her toes were shorter and her feet
were wider and squarer. They looked like Russian rather than
English feet.
- D. J. Morton: “Significant Characteristics of the Neanderthal Foot,” Nil,
Vol. 26, No. 3 ( 1926), pp. 310-4.
3 Piveteau, op. cit., p. 462.
548
The Caucasoids
The Height and Build of the Western Neanderthals
According to the Neanderthal legend, he was a squat,
stunted fellow, about five feet one inch tall, or 155 cm. As indi-
cated by careful calculations from his long bones, La Chapelle
aux Saints stood five feet four and a half inches tall, or 164 cm.,
about half an inch taller than the Frenchmen who lived in the
region of his cave at the time his remains were excavated. Nean-
derthal, Spy 2, and La Ferrassie 1 were of about the same height,
five feet four, five feet four, and five feet six ( 163 cm., 163 cm.,
and 165.7 cm.), whereas the female La Ferrassie 2 was four feet
ten inches, or 148 cm. tall, as might be expected from the rela-
tively small size of her head.
With large heads, deep chests, heavy bones, and large feet, the
western Neanderthals must have been heavy for their stature,
probably a good 160 pounds or more. They were prime examples
of what students of human constitutional types call mesomorphs.
They were indubitably muscular, but some of their muscles were
more developed than others. The muscles of the upper back and
neck had to be strong to support the weight of the head and par-
ticularly of the prowlike face. The muscles that roll the humerus
outward from the trunk were powerfully developed, but the biceps
and triceps of the arm need not have bulged greatly because of
special leverage. The forearm, however, must have been very im-
pressive. The calf muscles also were probably as filled out as those
of Alpine mountaineers. People built more or less like these Nean-
derthals may be seen today in the Abruzzi Mountains, in the Alps,
and in Bavaria. Whether the resemblance is due to the infiltration
and absorption of Neanderthal genes into later populations, or
merely to parallelism, we do not know.
The Fate of the Western Neanderthals
Without much doubt the Neanderthal population of western
Europe was greatly reduced by the end of the Wiirm I glacia-
tion, but it did not die out completely until after the Upper
^ ---
The Central European Neanderthals 549
Paleolithic people arrived during the Gottweig Interstadial. In
France the Mousterian culture lasted into the beginning of that
same warm period in at least one site, the Grotte du Loup at Arcy-
sur-Cure. Late Mousterian sites also exist in Italy and Spain. In a
Spanish site, Pinar, near Granada, in a level above that which
contained the Neanderthal frontal bone already mentioned, J-C.
Spahni found a basically Mousterian industry that contained
traces of Upper Paleolithic techniques of tool-making, implying
contact between Neanderthal and Upper Paleolithic peoples.4
The date of the level is Wiirm I— II, the Gottweig Interstadial. In
it Spahni also found a mandible of Upper Paleolithic type.
The implication of this and other contemporaneous sites is that
the two peoples met, and mixed. There is no valid anthropologi-
cal or biological reason for some of the western Neanderthals not
to have been absorbed into the immigrant populations. At least in
these southern regions some Neanderthal genes must have been
taken into the Upper Paleolithic pool. The Neanderthals became
extinct; of that there is no question. But their extinction was
probably of the usual human form, extinction by absorption.
Some of the physiological peculiarities of the Neanderthals prob-
ably became useful in the mixed population that followed, par-
ticularly with the advance of the second Wiirm ice sheet.
The Central European Neanderthals
Central Europe is as poor in Neanderthal remains as
western Europe is rich. The difference is not due to a lag in search
and excavations, for archaeologists in Central Europe have been
nearly as busy in the last century as their western colleagues.
Mousterian sites are rare in the middle of the continent. They are
concentrated in a few favored spots, like the Biikk Mountains of
north central Hungary. These are not really mountains but prom-
ontories on a small, detached highland no more than a thousand
feet in elevation. It was colder in central than in western Europe
4 It is also possible, but less likely, that Upper Paleolithic invaders found
Mousterian artifacts and reworked them into their own kinds of tools, which they
then left in the cave.
550
The Caucasoids
i]
J
during Wiirm I, just as it is today, and the difference in winter
temperatures between the Upper Danube and the Dordogne could
have been critical for the survival of any kind of people living at
that time, including Neanderthals.
The skeletal material which we have for that period is extremely
scanty. It consists primarily of three lower jaws and a total of
thirty-five teeth. In addition, we have some of the postcranial
bones of an undersized adult female, a child’s skull as thoroughly
smashed as Humpty-Dumpty, and one toe bone. That is all. They
are listed in Table 28.
The Three Mandibles
The mandible first found came from the Sipka Cave near
Stramberk, North Moravia, in 1880.5 Not until 1955 did Czecho-
slovakian archaeologists decide that it belonged to a Mousterian
culture, probably in Wiirm I. Destroyed in a fire in 1945, it was
the front part of the jaw of a child who had not yet cut his
permanent canines; his premolars were not only uncut but still
rootless. It was a large and stout jaw for a child of nine.8 The
sagittal profile, below the bulge of the dental area, ran straight up
and down, and the center of the lower border projected down-
ward, as in La Ferrassie 1.
The Ochoz mandible was discovered in a cave in Central
Moravia in 1905. At the time no implements were found with it.
Later, in 1954-1955, archaeologists discovered two successive in-
dustries, a Mousterian from Wiirm I and an Upper Paleolithic
from Wiirm II or III. It was impossible to tell which industry the
Ochoz jaw belonged to, but it has been assigned to the earlier
one on morphological grounds.7 The fauna fits both dates.
Although both ascending rami are missing, about 1 cm. of the
5 R. Virchow: “Der Kiefer aus der Sipkahohle und der Kiefer von La Naulette,”
ZFE, Vol. 14 ( 1882), pp. 277-310.
K. J. Maska: Der diluviale Mensch in Miihren ( Neutitschein, 1886).
E. Vlcek: “Die Reste des Neanderthalmenschen aus dem Gebiete der Tsche-
choslowakei,” NC, 1958, pp. 107-20.
6 Sagittal height = 30 mm.; thickness = 14. mm.
7 It is listed in the Catalogue des Hommes Fossiles, edited by Vallois and
Movius and published in 1952, as Wiirm II or III. The archaeological work was
done later. See Vlcek: op. cit.
55i
The Three Mandibles
body is left to the rear of the left third molar and in this portion
there is no trace of the beginning of the left ascending ramus that
would be there if it were an Upper Paleolithic type of jaw. This
is a specific Neanderthal condition. Also, the molars are tauro-
dont. Foi these reasons I consider the Wiirm I date correct.
The whole lower portion of the jaw is missing, which makes it
impossible to tell whether it had a chin, but the profile of the
symphyseal region is an open double curve. The jaw was slightly
prognathous, and probably had a slight chin, like some of the
Krapina mandibles.
The Subalyuk mandible was found, along with other human re-
mains, in a cave located near the edge of the Biikk Mountains in
Hungary. The cave was named after Michael Suba, a legendary
bandit who used to hide in it. After human remains had been
unearthed there, the Hungarian government renamed it the
Mussolini cave, and this name persists in the literature.8 It is the
only site in Hungary containing stratified Mousterian artifacts.
There were two cultural levels, a “high” Mousterian of fine quality,
and a later, decadent Mousterian. The human remains are said
to have come from the latter level.
The mandible is that of an adult woman. It consists of two dis-
connected pieces: the chin portion, extending from the left lat-
eral incisor to the right first premolar; and the left side of the
body from the second premolar to the third molar and beyond,
including a large part of the left ascending ramus. Morphologi-
cally it fits the western Neanderthal pattern except that it, like the
Ochoz jaw, is prognathous. As in La Ferrassie 1, the bone is very
thin.9 The ascending ramus slopes back widely, and the distance
from the rear edge of the third molar is as great as in La Ferrassie.
Also, the inside of the chin portion has the same exaggerated re-
lief for the insertion of the digastric muscles as does the French
jaw.
In general, these three mandibles are closely similar to those
SL. Bartucz, J. Dancza, F. Hollendonner, O. Kadic, M. MottI, V. Pataki,
E. Palosi, J. Szabo, and A. Vendl: “Die Mussolini Hohle (Subalyuk) bei
Cserepfalu, GHSP, Vol. 8, No. 14 (1939), pp. 1—320.
9 The symphysial height is 14 mm.; the height at the canine level, 33.7 mm.;
and its thickness, 13 mm., compared to a thickness of 16 mm. for La Ferrassie 1’
a male.
552
The Caucasoids
of the western Neanderthals except for one feature — a consider-
able prognathism. In this they favor their central European prede-
cessors, the men of Ehringsdorf and Krapina.
The teeth imbedded in these jaws are also Neanderthaloid. Al-
though all of them are within the modern size range, they are not
entirely modern in form. As far as I can tell, all three jaws, and
certainly Subalyuk, have taurodont molars. The unworn incisors
of Sipka are moderately shoveled.
The Postcranial Bones from Subalyuk
With the Subalyuk mandible were found other parts of the
woman’s skeleton: a fragmentary atlas (first cervical vertebra),
three dorsal vertebrae, one lumbar vertebra, one fragmentary
sacrum, one manubrium sterni (the lower section of the breast
bone), one metacarpal, one patella, three metatarsals, and two
toe phalanges. Together, these pieces indicate a small, poorly
muscled woman of the usual Neanderthal type. However, two,
the sacrum and the manubrium, are notable.
The sacrum consists of two pieces broken during excava-
tion, including the top. It is tiny and would fit an Andamanese
Pygmy, or a European child. Whether it really belongs with the
rest of the skeleton is an open question.1 The manubrium of the
sternum is a bone rarely found in fossil deposits, because it is soft
and spongy and decays rapidly. This is the only Neanderthal
manubrium we have. It is small, like the rest of the skeleton, but
it is also distinctive in form. On its inner side it is deeply concave,
with a depth of 7.5 mm. Most manubria are flat or slightly concave.
This can only mean that the woman had a deep chest, a standard
Neanderthal feature.
The Subalyuk Child’s Skeleton
With the woman was found the skeleton of a child; it was
badly smashed, mostly in excavation. Only the skull has been
1 Published photographs of this excavation show men swinging heavy pickaxes
over their heads.
Significance of Neanderthal Remains from Central Europe 553
studied. Bartucz painstakingly reconstructed the brain case from
about sixty pieces and the face from about twenty. The child was
about six or seven years old, judging from the "eruption of the
teeth, which have not been studied. It has been compared with
the eight-year-old La Quin a skull from France, and is similar in
all peitinent respects. There is no canine fossa, the nasal opening
is great for a child of that age, and the left nasal bone is concave
at the top and convex below, as befits a Neanderthal nose.
The Rumanian Neanderthal Toe Bone 2
Rumania’s contribution to the Neanderthal problem is so far
negligible. It consists of the discovery of one phalange from a
second toe, found in a rare Mousterian site in the Transylvanian
Alps, a region that ethnically and geographically belongs to
Hungary. Because no one whom I know can tell a single toe
phalange of a Neanderthal from that of an ordinary European,
this discovery only adds an osteological item to the archaeological
evidence that people of Mousterian culture lived in those moun-
tains in Wiirm I.
The Significance of the Neanderthal Remains
from Central Europe
The extremely limited roster of Neanderthal remains
from central Europe leads us to the following conclusions. Com-
pared to France, Italy, and Spain, central Europe was very
sparsely inhabited in Wiirm I times, probably because the cli-
mate was too cold. The people who lived there were Neander-
thals, but they differed from their western relatives in having a
considerable alveolar prognathism, a trait also seen in their prede-
cessors of the Last Interglacial. It is doubtful that they were
numerous enough, or in sufficient contact with their neighbors, to
have served as a genetic bridge between the western Neander-
“ M. N. Morosan: “Les Restes de I’Homme Fossile en Roumanie ” IGC 1936
pp. 1239-47-
554
The Caucasoids
thals and the contemporary inhabitants of the Soviet Union and
western Asia.
Yet they seem to have survived into the Gottweig Interstadial,
long enough to have made contact with the first wave of Upper
Paleolithic invaders. In a cave called Veternica, near the village
of Gorni Stenjevic in the neighborhood of Zagreb, Yugoslavia,
three skulls were found under a stone cover in a cultural level
attributed to the threshold between the Mousterian and the
Aurignacian.3 According to Malez, who found them, they are
youthful and modern in type. One level below, another skull
turned up in soil containing Mousterian implements. C. Loring
Brace, an American anthropologist who saw this skull in 1959,
says that it is an ordinary modern sapiens skull, associated with
an ordinary Mousterian industry, dated at the first interstadial
of the last glaciation.4 Croatia, then, vies with Spain as the place
where the two then existing kinds of Caucasoids, Neanderthal
and modern European, may have met.
Neanderthal Remains from the Soviet Union
From the Alps to the Himalayas, the northern zone of the
Palearctic region, west of the Tian-Shan Mountains, is separated
from its smaller and more southern portion by an east-west moun-
tain barrier, which was glaciated locally in several centers during
the Wiirm period. Even unglaciated, these mountains have always
been difficult to cross, and human traffic has usually been routed
to either side of them. That is why the Carpathians, Caucasus,
and Hindu Kush are refuge areas still inhabited by culturally
archaic peoples.
From the mouth of the Rhone to India there is only one major
gap in this mountain wall. It is the Bosporus gateway. Because
of this open passage, which was dry land during much of the
Pleistocene, the shores of the Black Sea were accessible to peoples
moving to and from the warmer lands of western Asia, notably
3 M. Malez: “Die Holile Veternica, eine neue palaolitische Fundstelle in Kroa-
tien, BSY, Vol. 3, No. 1 (1956), pp. 11-12.
4 Letter dated October 23, 1959.
555
The Kiik-Koba Tooth and Limb Bones
Palestine and Lebanon. The Acheulian hand axes found in
southern Russia owe their presence to penetration from the Levant
rather than from western Europe. In the days of the Neander-
thals the sea-level route between the Levant and the Black Sea
must have been easier to travel than the chill mountains and
forests of central Europe. The Crimea, a vacation spot today, was
a particular favorite of the Neanderthals. It contains many
Mousterian sites, two of which, Kiik-Koba and Starosel’e, have
yielded human remains.
The Kiik-Koba Tooth and Limb Bones 5
Kiik-Koba is a cave lying at an altitude of about 1,400 feet
m Zuya River Valley, about 15 miles east of Simferopol. It was
excavated in 1924 by Bonch-Osmolovskii, who found two cultural
levels: a relatively crude flake industry with a temperate fauna,
and above it an evolved Mousterian with a cooler fauna. In the
middle of the limestone floor of the cave the remains of a human
body were buried in a shallow trench. Like the French Neander-
thal burials, the trench ran east and west. At some point but still
within glacial times, the burial had been disturbed and all the
bones removed excepting the feet and left lower leg, which were
left in their original position. From the disturbed area parts of
the right hand and one incisor tooth were recovered. Although
this is not certain, the skeleton was probably associated with the
later culture, and belonged to Wiirm I. It might be older, but it
is not younger.
The incisor tooth was worn down to the neck. In size and form
it could fit in the western Neanderthal series.
The right hand is represented by the trapezoid and trapezium
(the wrist bones nearest the thumb), by the first and fourth
metacarpals, and by three proximal, four middle, and three
terminal phalanges. As indicated in Fig. 75, not one of the five
5 C. A. Bonch-Osmolovskii: “Paleolit Krima, No. II, 1941, Kist iskopaemogo
Cheloveka iz Grata Kiik-Koba” ( Moscow-Leningrad: Izdat. Akad. Nauk SSSR-
i94i)-
Bonch-Osmolovskii: “No. Ill, 1954, Skelet stori i goleni iskopaemogo Chelo-
veka iz Grata Kiik-Koba” (Moscow-Leningrad: Izdat. Akad. Naut. SSSR; 1954).
H. Ullrich: “Neanderthalfunde aus der Sowjetunion,” NC, 1958, pp. 72.-106.
556
The Caucasoids
Fig. 75 Neanderthal Hands and Feet. A. Hand skeleton from Kiik-Koba (after
Bonch-Osmolovskii, 1958); B. Foot skeleton from Kiik-Koba (after Bonch Os-
molovskii, 1958); C. Footprint in the cave of Basua, Toirano, Liguria, Italy (after
Leonardi, 1958 ) ; D. Sole of the right foot of an Alakaluf Indian woman, Welling-
ton Island, Chile (after a photograph by the author); E. Another footprint from the
cave of Basua (after Leonardi, 1958). The Neanderthal hand skeleton (A) is short
and broad, like those of many modern Russians; but no single finger is complete.
The foot skeleton (B) is of the same general build. Of the two Neanderthal foot-
prints, E is the longer ( ca . 20 cm.). Both show a gross similarity to that of the
Alakaluf woman, who walks barefoot every day in briny water. The resemblance
is not racial, but convergent and adaptive. Both Neanderthals and Fuegian Indians
are, or were, cold-adapted.
finger skeletons, including the metacarpal and all phalanges, is
complete; it is therefore impossible to reconstruct the whole hand
in its original proportions. The bones were nearly identical in size
with those of La Chapelle aux Saints and larger than those of La
Ferrassie 2, which suggests that Kiik-Koba was a male. The meta-
carpals and phalanges are very broad at the articulating ends,
and the hand itself must have been correspondingly wide. The
proximal end of the fist metacarpal ( thumb ) is rounded as in La
Chapelle.
The right foot, which is nearly complete, closely resembles that
557
The Infant Skeleton of StaroseVe
of the woman from La Ferrassie, except that it is larger. The total
length of the foot skeleton is about 226 mm., or nine inches. It was
undoubtedly masculine. The foot is long in the tarsal portion,
and short in the toe. The great toe is shorter than the second one.
All bones are broad, particularly in their articulating surfaces.
The measurements of the individual bones closely match those
of La Chapelle aux Saints. This foot could have made the prints
found by Baron Blanc in an Italian cave.
Its left tibia is almost exactly the same length as the left tibia
of La Chapelle aux Saints, and it is equally robust, but straighter.
The fibula that matches it is also a stout bone, and has a large
distal end where it articulates with the astragulus. It, too, is
straight. Kiik-Koba man was probably about as tall as his French
counterpart, and similarly built.
The Infant Skeleton of StaroseVe 6
I n 1952 a Russian archaeologist, A. A. Formosov, discovered the
skeleton of an infant in a cave in the Crimean village of Starosel’e,
overlooking the Tschuruk-su River. Associated with it were Late
Mousterian implements. The date was probably late Wiirm I.
Like most skeletons of infants that have been unearthed, this
one was badly squashed. But M. M. Gerasimov, a Russian sculp-
tor who is also an anthropologist, painstakingly restored the skull
(see Fig. 57). The result is a startingly modern skull; in fact it
looks like a caricature of Mr. Molotov, with his bulbous forehead
and square chin. The resemblance is superficial, however, because
Mr. Molotov is an adult and the Starosel’e infant was only
eighteen or nineteen months old.
To the unpracticed eye, the skull looks completely modern and
completely Caucasoid. But babies’ skulls are deceptive, and we
have no Neanderthal skulls of equal age to compare it with. The
forehead is high and steep, and somewhat bulbous. It is even
more strongly rounded and bowed forward than most modern
6 I. I. Roginskii: “Morfologischeskie Osobennosti Cherepa Rebenka i Pozdne-
must’erskogo Sloia Pschery Starosel’e,” SE, Vol. l (1954), pp. 27-39.
Ullrich: op. cit., pp. 72-106.
558
The Caucasoids
1
baby skulls of the same age. The face is shorter than most com-
parable baby faces; the mandible has a firm chin and the lower
borders of the mandible spread outward. It has a distinct canine
fossa. The back of the head is high and rounded. All these essen-
tial features are completely modern.
On the other hand, the vault is thick for an eighteen-months-
old baby, especially the lower part of the forehead. The mastoids
are weakly developed, and the milk incisors, although wide, lack
the thickness of Neanderthal milk incisors. The milk molars, how-
ever, are large. In Roginskii’s opinion, it would have developed
heavy brow ridges had it lived, but it had none of the prowlike
features of the Neanderthal nasal region.
It is either an early example of modern Caucasoid Homo sapi-
ens, a product of mixture with local Neanderthals, or the end re-
sult of an evolutionary progression from Neanderthal to mod-
ern European man. Which alternative is correct cannot be decided
at this point, but we shall come back to the question after describ-
ing the Wiirm I populations of the Levant.
The Youthful Neandeiihal of Teshik-Tash 7
I n 1938 a Russian archaeologist, A. P. Okladnikov, excavated a
Mousterian deposit in a cave located in the flank of a gorge in the
Baisun-Tau Mountains of southwestern Uzbekistan, about 78
miles south of Samarkand and 60 miles north of the Afghan bor-
der. The cave was Teshik-Tash. In it Okladnikov found five suc-
cessive layers of habitation deposits, with hearths, Mousterian
implements, and animal bones, 84 per cent of which were of one
7 A. P. Okladnikov: “Issledovani Musterskoi Stoianki Pogrebenia Neadertal’tsa
v Grote Teshik-Tash, Iuzhnyi Uzbekistan,” Sbornik Teshik-Tash (Moscow, 1949),
pp. 7-85.
N. A. Sinelnikov and M. A. Gremiatskii: “Kosti Skeleta Rebenka-Neandertal’tsa
iz Grota Teshik-Tash Iuzhnyi Uzbekistan,” ibid., pp. 123-36.
Gremiatskii: “Cherep Rebenka-Neandertal’tsa iz Grota Teshik-Tash Iuzhnyi
Uzbekistan,” ibid., pp. 137-82.
Weidenreich: “The Paleolithic Child from the Teshik-Tash Cave in Southern
Uzbekistan (Central Asia),” AJPA, Vol. 3, No. 2 (1945), pp. 151-62.
Movius: “The Mousterian Cave of Teshik-Tash, Southeastern Uzbekistan,
Central Asia,” ASPR, Vol. 17 ( 1953), pp. 11—71.
Ullrich: op. cit.
The Youthful Neanderthal of T eshik-T ash
species, Capra sihirica, a local wild goat. The other animal bones
were from the modern horse, leopard, bear, hyena, and small
rodents. All except the hyena, which was of the cave-dwelling
variety, exist in Uzbekistan today.
In a shallow grave under the top layer Okladnikov found the
remains of a nine-year-old boy surrounded by five pairs of wild
goat horns. He considered it a burial, although few of the post-
cranial bones were present and the skull was badly broken. Since
1938, some authors have expressed the belief that the burial had
been disturbed by a hyena, others that the bones had been stripped
of their flesh before being placed in the cave.8
Whichever may have been the case, of far greater importance
for our purposes is the determination of the age of the find. The
Russian scientists consider it to have been deposited in the Riss-
Wiirm Interglacial; and although Movius believes that its date is
most probably the Wiirm I— II Interstadial, he does not exclude
the possibility of a late Third Interglacial age. In favor of a
Wiirm I— II Interstadial dating are three facts: the climate was the
same as it is today; the fauna is modern; and the industry is an
evolved Mousterian. If this date is correct, the Teshik-Tash child
was roughly contemporaneous with an Upper Paleolithic blade
culture 150 miles to the south, in Afghanistan.9
Gerasimov skillfully reconstructed the child’s skull from more
than a hundred pieces, most of which are very small. However,
about forty are large enough to permit identification and the final
product is a nearly complete cranium. The mandible was intact.
The cranial capacity is 1,490 cc., and had the child lived to
maturity, this capacity probably would have reached a figure of
1,600 cc. Its basic dimensions, length, breadth, and basion-bregma
height ( 185 mm., 144 mm., and 132 mm. ) , can easily be matched
among skulls of modern children of both European and Mongoloid
populations, and its height is greater than that of the skull of any
8 The argument of the anti-hyena school is that when a hyena eats a femur,
he starts at the head of the bone and works down to the marrow cavity. In the
Teshik-Tash specimen the femur head was intact, except for the epiphysis, and
only the shaft was broken. See Ullrich: op. cit.
9 In the cave of Kara Kamar, near Haibak. See my The Seven Caves (New
York: Alfred A. Knopf; 1957). However, between the Upper Paleolithic blade cul-
ture and a Mesolithic level was a layer containing a still undiagnosed flake culture.
560
The Caucasoids
adult western European Neanderthal. The forehead is high and
well rounded, the occiput somewhat bun-shaped, and the lamb-
doid region somewhat flattened. The brow ridges are heavy for a
nine-year-old child, and continuous across the mid-line of the
skull. The orbits are high and the nasion depression slight. In all
pertinent details it closely resembles the frontal fragment and
nasal bones of the Pinar child from Spain, said to be about eight
years old.
The nasal opening is broad. There probably was no canine
fossa, although this is not certain because the parts of the maxillae
in which this feature is seen are missing on both sides. The
maxillae are not, apparently, very much inflated by sinuses, in the
western Neanderthal manner, and there is little prognathism
either nasal or alveolar. The upper index of facial flatness is 22, a
Caucasoid figure, and the upper edge of the orbit clearly over-
hangs the lower.
The mandible has a definite if weakly developed chin, and
looks square in front because the lower permanent canines are
fully formed but unerupted and imbedded in the bone. The
teeth are larger than those of most western Neanderthals, but
within the modern size range. The upper incisors are slightly
shoveled, and thick at the base of the crown, with basal shelves
typical of the Neanderthals.
In sum, this skull is difficult to evaluate because we lack ma-
terial of a comparable age with which to contrast it. Except for
the fragmentary Pinar specimen, all other youthful Neanderthal
crania are either younger or older. Although the Teshik-Tash skull
bears many of the hallmarks of the western Neanderthal, it is
more modern-looking than most if not all of the western Nean-
derthal skulls, because the vault of its brain case is higher and less
baggy-looking than theirs, and its face seems less muzzlelike and
less puffy in the maxillae.
It is the kind of skull one would expect to find in a Neander-
thal population which had not undergone the specializations of
the western Neanderthals, or which had lost them, either through
progressive evolution, mixture with a more modern Caucasoid
people, or both. Weidenreich, in 1945, saw in it certain features
characteristic of Sinanthropus, but to my mind these are less
The Eastern Neanderthals of Shanidar 561
marked in the Teshik-Tash child than in Krapina or the western
Neanderthals.
The postcranial bones include the atlas; twelve ribs; two clavi-
cles, one of which is broken; the left humerus, which lacks both
ends; the upper part of the right femur; the left tibia, which lacks
both ends, and the shafts of both fibulae. These have been de-
scribed by Sinelnikov and Gremiatskii.
The atlas is large, slenderly built, and of only medium height;
its opening for the medulla is large. The clavicles are of normal
size for a nine-year-old boy, and they are curved more than mod-
ern clavicles. The western Neanderthal clavicles are the opposite —
they are less curved than modern ones. The ribs are more curved
than those of the western Neanderthals, but some of them are tri-
angular in section, like one of the ribs of La Chapelle aux Saints.
The humerus shaft is straight and without torsion. The femur is
modern in size and proportions, with none of the extreme bowing
seen in western Neanderthal femora. Its neck goes off the shaft at
a wider angle ( 130° ) than that of the western Neanderthal ones.
The tibia is thick, triangular in section, and straight, and the
fibulas are straight and of the proper length for a modern Cauca-
soid.
From the neck down, therefore, the nine-year-old boy of Teshik-
Tash was a modern Caucasoid in all essential respects; his body
was even less Neanderthaloid than his skull. He was not, however,
unique. We shall see others like him in Western Asia; in Iraq,
Palestine, and Lebanon.
The Eastern Neanderthals of Shanidar
I n 1949 a Turkish archaeologist, I. K. Kokten, found two teeth
in a Mousterian deposit in a cave near Adalia, on the southern
shore of Anatolia. These teeth have been identified as Neander-
thaloid.1 In 1954 M. S. §enyiirek, a specialist on fossil teeth, and
E. Bostanci, a physical anthropologist, discovered three more such
teeth in a lake-shore cave at the foot of Musa Dagh, some 700
1 M. S. §enyiirek: “A Short Preliminary Report on the Two Fossil Teeth from
the Cave of Karain . . . ,” Belleten, Vol. 13, No. 52 (1949), pp. 833-6.
562 The Caucasoids
miles to the east-northeast.2 These widely separated sites indicate
the presence of Neanderthal man, and his Mousterian culture, in
Turkey.
In 1949 I found a piece of strongly bowed ulna and a lower
incisor tooth in Bisitun Cave in the western flank of the Zagros
Mountains of Iran, along with an evolved Mousterian industry,
and also what seems to be a small piece of human femur shaft in
Tamtama Cave, on the east side of the Zagros, in Iranian Azerbai-
jan.3 I am not certain that these three specimens belonged to
Neanderthals, although the ulna fragment looks as if it did; all
I know is that some kind of man lived in or near the Zagros Moun-
tains during Wiirm I or the Wiirm I— II Interstadial. The Turkish
teeth and these Iranian finds half encircle the site of Shanidar, in
which Ralph Solecki found seven Neanderthal skeletons between
1953 and i960.
Shanidar is a huge, majestic cave in the western Zagros of north-
ern Iraq, inhabited from early Mousterian times to the present;
several Kurdish families still live in it. Now and then, owing to an
earthquake or to the formation of ice in cracks in the roof, slabs
of limestone crash to the floor, killing everyone below. In 1953
Solecki found a baby’s skeleton; in 1957, three adult skeletons,
numbered 1, 2, and 3; and in i960 three more.
All lay in Mousterian deposits. Shanidar 1 has been given a
Carbon-14 date of 46,000 ± 1,500 years; 4 Shanidar 3 was perhaps
a few hundred years older. Shanidar 2 and the baby were prob-
ably about 60,000 years old, and the i960 skeletons probably
60,000 years old or older. Shanidar 1 was found at the very top of
the Mousterian deposit, indicating that the cave may have been
abandoned during the height of the Wiirm I cold, after which,
judging by the overlying deposits, it was reoccupied by Upper
Paleolithic people about 35,000 years ago, during the Wiirm I— II
Interstadial.
2 Senyiirek and E. Bostanci: “The Excavation of a Cave Near the Village of
Magraeik in the Vilayet of the Hatay, Preliminary Notice,” Anatolia, Vol. l
(1956), pp. 81-3.
3 C. S. Coon: “Cave Explorations in Iran, 1949,” VMM, 1951.
4 R. S. Solecki: Three Adult Neanderthal Skeletons from Shanidar Cave,
Northern Iraq,” SRP, No. 4414 (1959-60), pp. 603-35. The C -14 date number
is GRO-2527.
I
ft
The Eastern Neanderthals of Shanidar 563
To date, the baby’s teeth have been studied,5 and called Nean-
derthal, and preliminary reports on Shanidar 1 have been pub-
lished by T. D. Stewart.5 Other reports should appear shortly.
Shanidar 1 and 2 were males, and §enyiirek believes that the
baby was a girl. The others have not yet been sexed. Shanidar 3
was not killed by rockfall but was buried against the cave wall.
A projectile point, found in his rib cage, may have been the cause
of death.
Shanidar 1 died at about age forty. He was approximately five
feet seven or eight inches tall ( 170-173 cm.), four or five inches
taller than the French Neanderthals whose statures have been
computed.' Judging from published photographs, his limb propor-
tions were normal for Caucasoids and unlike those of La Chapelle
aux Saints and his western European companions.
Because Shanidar 1 had been born with a damaged brachial
plexus, his right scapula and clavicle were atrophied and his
right arm hung limply until some time before death, when a Nean-
derthal surgeon amputated it above the elbow, presumably with a
flint knife. Either before or after this successful operation, Shani-
dar 1 had been severely wounded by blows with a sharp instru-
ment around and particularly above the left eye, which may then
have been blinded. He also had a bone lesion from a blow on the
right parietal. Despite these injuries and adventures he died at
home, standing in his own cave, crushed by a slab of limestone.
When the complete measurements are available,8 Shanidar l’s
skull may prove to be the largest fossil-man skull of its date or
earlier yet found. The cranial capacity is probably well over
5 Senyiirek: “The Skeleton of the Fossil Infant Found in Shanidar Cave,
Northern Iraq, Preliminary Report,” Anatolia, Vol. 2 (1957), pp. 49-55.
Senyiirek: “A Further Note on the Paleolithic Shanidar Infant,” Anatolia, Vol.
2 ( 1957), PP- 111-21.
6 T. D. Stewart: “First Views of the Restored Shanidar I Skull,” Sumer, Vol.
14, Nos. 1-2 ( 1958)* PP- 90-6.
Stewart: “Restoration and Study of the Shanidar I Neanderthal Skeleton in
Baghdad, Iraq,” YAPS, 1958, pp. 274-8.
Stewart: “The Restored Shanidar I Skull,” SRP, No. 4369 (1958), pp. 473-8.
Stewart: “Form of the Pubic Bone in Neanderthal Man,” Science, Vol. 131,
No. 3411 (i960), pp. 1437-8.
7 This figure was calculated by Stewart from the length of the ulna; a more
accurate estimate will be made when the leg bones are described.
8 Any figures given here are tentative, made from scale drawings and photo-
graphs.
564
The Caucasoids
1,700 cc. The forehead is sloping, but the parietal arc is greater
than the frontal, as in modern men, and both the parietal and
occipital profiles are well rounded. Seen from the front and rear,
the brain case lacks the bagging or “soft watch” appearance of
the western Neanderthals; in this respect it resembles Teshik-
Tash.
The face is as long as those of La Chapelle aux Saints and La
Ferrassie 1; and characteristically of Neanderthals, the bizygo-
matic face breadth is less than the breadth of the brain case. The
brow ridges are heavy and overlie extensive sinuses; but unlike
those of the western Neanderthals, they do not form a continuous
bar over the nose but are divided, as in most other early skulls.
The nasal bones, which are intact, rise high under the frontal,
and the nasal profile is projecting. The orbits, which are large
and rounded, fall away to either side, giving the skull an index
of upper facial flatness of about 27, which is almost super-
Caucasoid.
Yet morphologically the face closely resembles those of the
French Neanderthals. The maxillary bone is puffy both under
the orbits and over the canines and does not have a canine fossa.
The face is prognathous only in the upper or nasal portion. In-
side the nasal aperture the floor of the cavity falls away steeply,
giving the nasal passages more depth than one would expect from
the size of the opening. La Chapelle aux Saints and La Ferrassie 1
apparently also possessed this feature, but one cannot be com-
pletely sure because of the damage to their nasal bones after
death.
The mandible is deep, and it is shaped essentially like those of
the French Neanderthals, except that it has more chin than all but
La Ferrassie 1, and its lower border is square in front. All the teeth
but the two lower median incisors are present, and all are heavily
worn. In size and form they resemble those of other Neanderthals.
The canines and incisors are worn not only on the crowns but
on their anterior surfaces, as if Shanidar 1 had held objects in his
teeth to compensate for the loss of his right hand. The teeth of La
Ferrassie 1 are similarly worn, and he also had a damaged arm.
Other details of the postcranial skeleton remain to be described.
However, Stewart has studied the pubic bones of Shanidar 1 and
The Inhabitants of Palestine During W iirm I 565
3. In each of them the upper ramus of the bone is thin and plate-
like, flattened from above and below. In modem Caucasoid
pelves, if not in all others, the ascending ramus of the pubis is
much thicker. No pubic bones of the western Neanderthals are
available for comparison, but, as we shall See presently, the pu-
bic bone of a Neanderthal woman from Palestine, Tabun 1, is simi-
lar to those of Shanidar 1 and 3. Stewart sees in this feature as
great a difference between the Neanderthals and modern men as
in the form of their head and face.
He has also stated that because Shanidar 1 and 3 lived late
in Wiirm I in what has always been a marginal, refuge area, they
lived too long to have sired modern Caucasoid man. But evolution
was moving at a faster pace on the eastern shore of the Mediter-
ranean, in Palestine particularly, as we have known since the ex-
cavation of the Mount Carmel caves some thirty years ago.
The Inhabitants of Palestine During Wiirm I
Although human remains from the Last Interglacial are
virtually nonexistent in Palestine, remains from Wiirm I are fairly
abundant. They come from six caves: Zuttiya, near the Sea of
Galilee; 9 Tabun and Skhul at Mount Carmel; 1 Jebel Qafza, near
Nazareth; 2 Shukba in the Wadi Natuf, seventeen and half
miles northwest of Jerusalem; and Amud, near Lake Tiberias.3
9F. Turville-Petre: Researches in Prehistoric Galilee, 1925-6 (London: British
School of Archaeology in Jerusalem; 1927).
Hrdlicka: op. cit.
1 A. Keith and T. D. McCown: The Stone Age of Mt. Carmel (Oxford: Claren-
don Press; 1939), Vol. 2.
C. E. Snow: “The Ancient Palestinian: Skhul V Reconstruction,” BASP, Vol.
17 (i955), pp- 5-10.
2R. Neuville: “Le Paleolithique du Desert de Judee,” A1PH, Mem. 24 (1951),
pp. 179-84.
3 Keith: New Discoveries Relating to the Antiquity of Man (London: Wil-
liams & Norgate; 1931 ), Chap. 13-14, for Shukba.
H. Suzuki, personal communication, Nov. 16, 1961, concerning the Amud Cave
skeleton.
For the entire group, see Howell: “Upper Pleistocene Men of the Southwest
Asian Mousterian,” NC, 1958, pp. 185-98.
The names of these sites are rendered here in a close approximation of correct
Arabic transliteration. Elsewhere the reader will see some of them spelled in
French transliteration, e.g., Djebel Kafzeh, which is incorrect. I have omitted
the words Mugharet al ( Cave of the ) wherever they occur.
566
The Caucasoids
Some of these remains may be older than the Shanidar skele-
tons, others of the same age or younger. It is difficult to tell be-
cause all but the new skeleton from Lake Tiberias, which has not
yet been studied, were excavated before the Carbon-14 dating
technique had been invented. It is not correct to call these skele-
tons, as a group, Neanderthals. Only one deserves that name in
the strict sense. Others are modern Caucasoids, and still others
intermediate between both extremes. One of them does not seem
to be either Neanderthal or modern Caucasoid, but looks Austra-
loid. This is a varied and complex group of skeletons which we
must use every means at our disposal to study, for in it may be the
key to the problem of modern European and western Asiatic Cau-
casoid origins.
The Galilee skull, from Zuttiya, consists of a frontal bone and
parts of the right zygomatic, the nasal bones, and the sphenoid.
Galilee man died at about twenty-five years of age. Keith first
called it a female, then a male; Hrdlicka identified it as an “effemi-
nate” male. From Tabun come a previously mentioned molar and
piece of femur shaft of Third Interglacial date, a complete female
skeleton known as Tabun 1, and a mandible, Tabun 2. Breccias
at Skhul have yielded parts of ten skeletons, consisting of five
adult males, Nos. 3, 4, 5, 6, and 9; two adult females. Nos.
2 and 7; one male child, No. 8; and two infants, Nos. 1 and 10;
and assorted postcranial bones of at least two other individuals.
Jebel Qafza’s contribution is five adults and one infant, all still
unstudied, and Shukba’s is seven children and one adult, also un-
described.4 So is the Lake Tiberias skeleton, discovered by H. Su-
zuki in 1961.
The date of the Galilee skull is uncertain. It is either the end
of the Last Interglacial or early Wiirm I. Tabun 1 and 2 are early
Wiirm I, and the rest are later Wiirm I. Recent investigations by
D. Brothwell and Z. S. Higgs at Cambridge 5 have shown that
Tabun 1 and 2 are about 10,000 years older than the Skhul group.
According to Brothwell, we can divide all these Palestinian
4 The Jebel Qafza remains were found in 1933; those of Shukba in 1928.
5 Z. S. Higgs and D. R. Brothwell: “North Africa and Mount Carmel: Recent
Developments,” Man, Vol. 61, No. 166 ( 1961), pp. 138-9.
Brothwell: “The People of Mt. Carmel,” PPS, October-December 1961, pp.
155-9-
Tabun and Galilee
567
skeletons into two lots, an older one, including Galilee and the
Tabuns, and a younger one, including those from Skhul, Jebel
Qafza, and Shukba.
Tabun and Galilee
The Tabun material from the Acheulian deposit of Last Inter-
glacial age consists of a femur shaft with both ends missing, and a
lower first molar tooth. The femur shaft is flattened from front to
back and has a weak linea aspera and no pilaster. The bone is not
strongly bowed. The tooth, which is badly worn, is indistinguish-
able from those of Tabun 1 and 2. Both femur and tooth can be
called Neanderthaloid, to the same extent that the specimens from
Ehringsdorf and Krapina can be so called.
The same may be said of the Galilee skull fragment. It has
heavy brow ridges divided in the middle by a depression, and a
well-rounded forehead of moderate breadth. The zygomatic
(malar) bone is not massive and is so shaped that the face may
have had a canine fossa. The orbits are of moderate size and
rectangular; the interorbital distance is great; and the nasal bones
are arched from side to side. The Galilee skull could have evolved
from a Last Interglacial pre-Neanderthal population like that of
central Europe.
Tabun 1 was a short woman, about five feet tall ( 154 cm. ) . Her
cranial capacity was 1,270 cc., the same as that of the female
Gibraltar 1, and the skull as a whole is small. The brow ridges are
heavy and continuous, the forehead retreating but curved, and
the parietal and occipital bones well rounded. There is no lamb-
doid flattening, nor an occipital bun. Although the vault is low,
with its greatest breadth well to the rear of the earholes, it lacks
the baggy configuration of the western Neanderthals. The orbits
are rounded; the face is long but in no sense prognathous; the nose
was apparently prominent and the chin retreating. The tooth line
forms an angle to the eye-ear plane, so that the mouth opened
somewhat downward; and the chinless profile of the lower jaw is
retreating to a large degree. The teeth are of moderate size for a
Neanderthal, but the incisors are characteristically thick, with a
distinct heel or shelf at the base on the tongue side.
The Caucasoids
568
The male mandible, Tabun 2, is large, deep, and squarish in
front. Its ascending ramuses are spread far apart to accommodate
a broad cranial base. In sagittal profile it is concave below the
tooth line and convex toward the chin, like the mandibles of
Ehringsdorf and Shanidar 1.
In general morphology the Tabun 1 skull and the Tabun 2 jaw
fall between the central Europeans of the Last Interglacial and
the western Neanderthals, and possess some special features that
are also present in the Skhul series. They also resemble Shani-
dar 1 in those features in which they deviate from the western
Neanderthals.
Tabun l’s postcranial skeleton confirms this diagnosis. Her
vertebrae, few of which were preserved, are short-bodied; her ribs
rounded in section and deeply curved; her sternum long. Her
scapulae resemble those of La Chapelle aux Saints, and her
clavicle is long for her stature. Her humerus is not stout, but its
head is directed somewhat upward as in the western Neander-
thals and its distal end is large, with a large olecranon fossa. Her
lower arm bones, radius and ulna, are widely separated as a result
of bowing; the head of the ulna is long, and the distal end of the
radius is wide. The hand is slenderer than that of La Ferrassie 1;
her thumb is short and her terminal phalanges long and slender.
Her pubic bone is flattened like those of Shanidar 1 and 3. Her
femur is flattened, but not very curved, and its neck is long. The
tibia is short and thick; the fibula rounded in section, like that of
La Chapelle aux Saints. Her foot is as long as that of a modern
western European woman of the same stature, but broader, with
the emphasis on the outer side of the foot; and her big toe was
short. In general, her feet resemble those of La Ferrassie 2 and
are less spatulate than Kiik-Koba’s.
Tabun 1 looks on the whole like a Third Interglacial European
woman who had acquired some but not all of the western Nean-
derthal specializations, or she might be a Neanderthal who had
lost some of these specializations through mixture. She fits into
the Asiatic Neanderthal population as we are beginning to know
it from Shanidar and Teshik-Tash; and her upper face and frontal
bone were a little more Neanderthaloid than the Galilee frag-
ment. Whether or not she was an ancestress of the tall men who,
The Skhul Skulls: No. 4 and His Group 569
some ten thousand years later, were buried in a neighboring cave,
we shall do our best to discover.
The Skhul Skulls: No. 4 and His Group
The skulls from the Skhul cave fall naturally into two
groups. Number 4 is the best preserved of the first group, and
Numbers 2, 7, and 9, although more fragmentary, are whole
enough to show a family resemblance to it. These four skulls show
both Neanderthal and modem Caucasoid features. The second
group consists of Skhul 5, the best known and most completely
restored of all, and probably also of Number 6, which is too frag-
mentary to be aligned with certainty in either camp. These skulls
are neither Neanderthaloid nor Caucasoid but belong to a dif-
ferent racial fine. The other four are either the skulls of babies, or
adult specimens too fragmentary for racial identification. As
Skhul 5 is generally considered the type specimen of the entire
cave population, the difference between the two groups may come
as a surprise to those who have not studied the other skulls with
equal care.
Skhul 4 was a male, about forty-five years old and five feet eight
and a half inches tall ( 174 cm. ) . Most of his skull is preserved, but
the region around glabella and nasion is missing. It is a large
skull, with a cranial capacity of 1,554 cc-> l°ng and low-vaulted,
but no lower than many modem European crania. It has a thick,
wide brow ridge, as big as that of La Chapelle aux Saints; a wide
mandible; a long face; and a deep palate.
In sagittal profile it looks much like La Chapelle aux Saints,
with two principal exceptions. Its occiput is rounded and not
bun-shaped. Its nose projected in a beak, but the facial skeleton
between the lower border of the nasal opening and the tooth line
is drawn backward and flat. There is no prognathism, either nasal
or alveolar. There is no canine fossa. The face is as long as those
of the western Neanderthals, and the palate as deep; but the
mandible has a firm chin. The parietals do not project sidewise as
in the western Neanderthals, and the zygomatic arches flare
widely, as in modern men who chew heavily, making heavy use
of their temporal muscles.
57°
The Caucasoids
If we shift our comparison from the western Neanderthals to
Shanidar i, we see that to make Shanidar I into Skhul 4 one need
only deflate the facial sinuses and pull back the palate. Skhul 4
looks like an evolutionary product from an earlier eastern Nean-
derthal base through Tabun toward a rugged, long-headed mod-
ern Caucasoid. Skulls which are called Nordic and which resem-
ble it in general form but with smaller brow ridges appear in
northern Europe from the Neolithic onward, and particularly in
the Iron Age.
Skhul 9, another male, had a cranial capacity of 1,587 cc., the
same type of brain case as Skhul 4, and the same hyperorthogna-
thous (the opposite of prognathous) facial structure, and prob-
ably the same kind of nose.
Skhul 2, a female five feet four inches tall ( 162.5 cm.), with a
cranial capacity of 1,300 cc., is a feminine version of Skhul 4. So,
as far as we can tell, for the skull is very crushed, was Skhul 7, a
female five feet two inches tall ( 158 cm. ) , with heavy brow ridges
and a long face.
The skulls of this group show an orderly progression from a
Neanderthal to a modem European form.
The Skull of Skhul 5
Except possibly for Skhul 6, a thirty-five-year-old male who
was five feet seven and a half inches tall (171 cm.) and of
whose skull we have only an occiput, Number 5 is unique. He
bears little resemblance to the Neanderthals before or the Nordics
after him; rather, he looks in many ways non-Caucasoid.
He was about thirty-five years old, five feet eleven inches tall
(180.6 cm.), and had a cranial capacity of 1,518 cc. The sagittal
profile of his skull starts with the usual heavy brow ridge, then
rises steeply and is well rounded the rest of the way, except for a
short interval of flattening at lambda. The skull has an ill-filled
look, like those of Australian aborigines, with prominent parietal
bosses. Seen from above, the brow ridge is nearly a straight bar,
instead of a bow curved backward at each end as it is in the Nean-
derthals and Skhul 4. Deep under glabella the stubs of his nasal
The Mount Carmel Teeth
57i
bones remain, fused into a single flat plate. His orbits are low and
square, and their sides form only shallow curves. This feature,
combined with his deeply set nasal root and flat nasal bones, give
him a flat upper face, with an index of upper facial flatness of
only 13.5, below any living racial mean and closest to some of the
ancient Australoid skulls, notably Wadjak 1, and to some of the
Bushmen. His upper jaw is very prognathous subnasally, and the
teeth in his lower jaw project well beyond his chin, as in Wad-
jak 2.
To my mind, Skhul 5 was not fully Caucasoid, and the features
in which he deviated from the Caucasoid line cannot be brushed
away as evolutionary grades. They are essentially Australoid.
These comparisons are not surprising in view of the geographical
position of Palestine at the crossroads of the Old World and the
fact that its fauna during Wiirm I was a mixture of Oriental,
Ethiopian, and Palearctic species.
Skhul 4 and his group seem to represent a station on an evolu-
tionary line from something like Swanscombe and Steinheim via a
local equivalent of the European Third Interglacial group to
Tabun 1 and an ancestor of Shanidar 1. Skhul 5 patently repre-
sents the product of a contact between the line just mentioned
and members of another subspecies.
The Mount Carmel Teeth
The picture that is beginning to emerge is supported by a
study of the teeth. Keith and McCown published descriptions of
sixty-five permanent and three milk teeth from Tabun and seven ty-
one permanent and twelve milk teeth from Skhul. None is mark-
edly taurodont, nor do any possess unusual enamel patterns like
those seen at Krapina.
In the unworn Tabun incisors and canines may be seen a mod-
erate shoveling with a labial heel, which takes the form of a
cingulum in one canine. One of the Tabun upper median incisors
has three tubercles and others have vertical ridging. The pre-
molars are relatively small, and the molars show no distinctive
features.
The Skhul teeth resemble those of Tabun but have less shovel-
572
The Caucasoids
ing and ridging. Keith and McCown saw in this series a se-
quence from Krapina to Tabun to Skhul. Yet between Skhul 4 and
Skhul 5 certain differences may be detected. Skhul 5’s palate is
enormous, big enough to fit Heidelberg or Wadjak 2. Its upper
incisors and canines show none of the Krapina-Neanderthal char-
acters; and in Flower’s index, the ratio between cheek teeth (the
premolar-molar row) and the basion-nasion diameter, it reaches
the high figure of 49.2 (see Chapter 8, page 353). This is ex-
tremely macrodont, beyond even the Tasmanian mean, whereas
Skhul 4’s Flower’s index is only 41.5, which is microdont and
Caucasoid. Although these contrasting ratios do not involve tooth
size as such, they involve the relationship between tooth size and
the proportions of the skull base and face. The combination is
racially diagnostic, and in this comparison its meaning is clear.
The Postcranial Skeletons of the Skhul Population
The postcranial skeletons of the Mount Carmel popula-
tion are not divided into the same three categories as the skulls,
but possess a separate dichotomy of their own. Essentially, Ta-
bun 1 and Skhul 7 form one distinct category, and all the others
form a second category. Skhul 4 and Skhul 5, which differ crani-
ally, are alike in the sense that both were tall men with slender
limb bones and relatively long forearms and lower legs. But
Skhul 7, a female whose skull belongs to the group exemplified
by Skhul 4, differs from the others found in the same cave in that
her long bones resemble, in certain respects, those of Tabun 1
and of the Neanderthals from western Europe and Shanidar.
We have vertebrae from Skhul 4 and 5 only. Skhul 5’s neck
was short, and its vertebrae small. Skhul 4’s dorsal vertebrae
were large enough to match his limb bones, but narrow from front
to back and perforated by large neural canals. His ribs were also
modern, except that the lower part of the rib cage was unusually
large. In both specimens the scapulae are small, indicating nar-
row shoulders, and they have a lipped groove along the axillary
margin which Boule had found earlier in La Chapelle aux Saints.
The clavicles are modern, the humeri long and slender, with small
olecranon fossae. The radii and ulnae are long in proportion to
The Meaning of the Mount Carmel Skeletons 573
their humeri, as in living Australoids and Negroids, but not long
enough to exceed the modern European range. The forearm bones
are curved, in the usual Neanderthal fashion, in only one skele-
ton, the female Skhul 7. The olecranon process, long in Neander-
thals, is of normal European proportions in the Skhul males.
Skhul 4 had large, long hands, and Skhul 5 had similarly shaped
hands, which were small for his stature. The pelves are modern,
without the pubic specialization of Tabun 1 and Shanidar; and
like the pelves of Krapina, they are narrow-hipped. The femurs
are long and straight except for that of the female Skhul 9, which
is very bowed. In the angle between the axis of the neck and
head of the femur to the axis of its shaft, Skhul 4 and 5 again part
company. Skhul 4 has an angle of 1220, a Neanderthaloid feature;
Skhul 5’s angle is 1320, a modern one. In this angle the faceless
Skhul 6 resembles Skhul 5 (its angle is 1350), as it does in other
respects.
The Skhul tibiae are long, so long in the males as to give the
two segments of the leg — the thigh and lower leg — Negroid or
Australoid proportions. All but that of Skhul 7 are sharp on the
leading edge, like modern European shins, but the tibiae of Skhul
7 are rounded, in Neanderthal fashion. We have only two whole
feet, from Skhul 4 and Skhul 7. Number 4’s foot is long, slender,
and modern; Number 7’s is similar to that of Tabun 1.
In sum, Skhul 4 and Skhul 5 are alike from the neck down, but
of the two, Skhul 4 was the stockier and had two Neanderthaloid
features, a large lower rib cage and a low femoral neck-to-shaft
angle. Skhul 5 lacked at least the second of these, and was more
Negroid or Australoid in build than Skhul 4. The female Skhul 7
shows a number of Neanderthaloid features in her extremities:
widely bowed forearm bones, a round-sectioned tibia, and a
Tabun-like foot. Skhul 9, for whom few bones are available, also
had a Neanderthaloid femur.
The Meaning of the Mount Carmel Skeletons
Ever since the discovery of the Mount Carmel skeletons,
anthropologists and anatomists, myself included, have been dis-
cussing their significance. In 1939 I expressed the opinion, later
574
The Caucasoids
shared by others,6 that the Mount Carmel population was the
product of a mixture between a local Neanderthal group com-
parable to those known from western Europe, and a more modern
stock. At the time I wrote, no eastern Neanderthals had been
described (although Teshik-Tash had already been discovered)
and Keith’s and McCown’s monumental work was not yet avail-
able. All we had to work with was essentially a preliminary de-
scription of Tabun 1, Skhul 5, and the western Neanderthal ma-
terial.
My present position is that, except for Skhul 5, the Mount
Carmel population shows an orderly descent from a local, Last
Interglacial population similar to that of Ehringsdorf and Kra-
pina, in a modern, Caucasoid direction. Like the Ehringsdorf-
Krapina group, this local population had become partially spe-
cialized in a Neanderthal direction and this specialization had
reached a peak in Tabun 1. In the Skhul skeletons this specializa-
tion was, apparently, being progressively lost. Whether as a sta-
tistical accident, for the series is small, or by some mechanism such
as sex-linkage, it was, apparently, being lost more rapidly in the
males than in the females.
Had Skhul 5 never been found, there would have been little
reason to talk of hybridization. But Skhul 5 was not only found;
it was publicized as the type specimen of the Mount Carmel
population. Its differences from the others buried in the same
cave at presumably the same period are not differences of grade
but of line. Its face is Australoid, and it resembles the skulls from
Wadjak more than any others that I have been able to find.
Strongly implied is a contact between Caucasoids and Australoids
in some part of southern Asia.
When other Palestinian skeletons of Wiirm I, from Jebel Qafza,
Shukba, and other sites yet to be excavated, have been described,
we shall be in a better position to interpret the racial variations
6 C. S. Coon: The Races of Europe (New York: The Macmillan Company;
1939), P- 38.
M. F. Ashley-Montagu’s review of A. Keith and T. D. McCown: The Stone
Age of Mt. Carmel, in AA, Vol. 42 ( 1940), pp. 518-22.
T. Dobzhansky: “On Species and Races in Fossil Man,” AJPA, Vol. 2 (1944),
pp. 251-65.
A. Thoma: “Metissage ou Transformation? Essai sur les Hommes Fossiles de
Palestine, L’Anth., Vol. 62, No. 1-2 (1958), pp. 30-52.
575
More About Neanderthal Origins
seen at Skhul. One Jebel Qafza skull, No. 6, 7 looks more modern
than any of the Skhul specimens. Essentially it is the same as the
Upper Paleolithic skulls from western Europe, which we shall
presently describe.
Egbert, the Boy from Ksar ‘Akil
I n 1938 two American Jesuit priests, Fathers J. G. Doherty and
J. Franklin Ewing, who were excavating the cave site of Ksar ‘Akil,
about seven miles northeast of Beirut, in Lebanon, discovered the
skeleton of a male child who had died in his seventh year. Later
Father Ewing restored the skull, and a plaster cast of it is avail-
able, but full details have not been published.8 Father Ewing
named it Egbert, and this name has had wide circulation.
The date is Wiirm I, the same as that of the Palestinian speci-
mens just described. Although the skull was broken in many
pieces and there are gaps in the reconstruction, there can be no
doubt about Egbert’s evolutionary and racial status. The brain
case is perfectly modern; there are no brow ridges; the forehead
is steep, the face orthognathous, and the chin firm. Egbert was a
modern Caucasoid, and probably would have grown into a man
resembling Jebel Qafza 6 had he lived. Moreover, the Starosel’e
infant might have grown to look like Egbert had his life been
spared for another six years.
Between the Wiirm I Palestinians and Egbert and Starosel’e,
there is a perfectly valid transition from earlier Caucasoid people
to Upper Paleolithic Europeans.
More About Neanderthal Origins
I n t h e light of what we have learned about the eastern Nean-
derthals, including in a wide sense the Palestinian and Lebanese
skeletons of Wiirm I, we may return to the discussion of Neander-
thal origins left open on page 558.
7 Judging from its photograph (see Plate XXVIII). Its dimensions have not yet
been published.
8 J. F. Ewing: “Human Types and Prehistoric Cultures at Ksar ‘Akil, Leba-
non,” F ICA, i960, pp. 535-9.
D. A. Hooijer: “The Fossil Vertebrates of Ksar ‘Akil, a Paleolithic Rock Shelter
in the Lebanon,” ZV, No. 49 ( 1961).
576
The Caucasoids
It is clear that the Neanderthals, eastern and western, were
derived, at least in large part, from the preceding Caucasoids of
the Last Interglacial, but it is not certain whether the distinctive
Neanderthal traits, both cranial and postcranial, arose through
mutation and selection alone, or were introduced into Europe
and western Asia by mixture with a non-Caucasoid population.
The finds from Teshik-Tash and Shanidar do not support, nor
do they completely disprove, the theory of mixture with Sinan-
thropus-descended Mongoloids across the mountain spine of north-
central Asia. However, the theory that the Saccopastore people,
whose skulls were the first to bear a Neanderthaloid stamp, could
have been the product of mixture between local Caucasoids and
North Africans is enhanced, purely fortuitously, by Keith’s and
McCown’s painstaking work on the Mount Carmel skeletons.
In studying the postcranial bones of this series, Keith and
McCown compared them with the skeleton of a South African
Bushman and expressed their surprise at finding in the latter
many striking resemblances to the skeleton of Tabun 1, and also
to that of the western Neanderthal, La Chapelle aux Saints. These
resemblances may be seen in the vertebrae, the ischial part of the
pelvis, the sciatic notch, and the limb bones, including the hands
and feet, and particularly the wrists and ankles.
These resemblances are too numerous and too striking to be dis-
missed as concidental on the grounds that it is a long way from
South Africa to Sicily. As I shall indicate in the next chapter, the
ancestors of the Bushmen, who were then full-sized people, prob-
ably lived in North Africa at the time of the Saccopastores, and
what remains we have of these pre-Wurm North Africans re-
semble Sinanthropus in details of the face, jaws, and teeth.
It is therefore easier to suppose that, if the Sinanthropus-like
features of the Saccopastores were due to race mixture, the alien
element came overseas from Cape Bon to Sicily, a distance of only
90 miles (60 if the immigrants stopped on the way at Pantellaria ) ,
rather than that they walked overland all the way from China.
Once they were in western Europe, and once the cold of Wiirm I
had set in, natural selection may have placed a premium on these
features, and the Neanderthal race came into being.
Of the three theories of Neanderthal origins — (1) mutation
The Upper Paleolithic People and Their Culture 577
and natural selection within a western European population dur-
ing the Last Interglacial and Wiirm I, (2) an infusion of favor-
able genes from the descendants of Sinanthropus in China, and
(3) a penetration of Sinanthropus-like genes from North Africa —
the third seems the most likely at the time of writing, but the other
two should not be forgotten. Ten years from now all three theories
may have been proven wrong.
The Upper Paleolithic People and Their Culture
In Europe the Wiirm I— II or Gottweig Interstadial, lasting
from about 40,000 to about 29,000 b.c., was a period of mild, but
not hot, climate, like that of the present, and of favored spots like
Palestine during Wiirm I. It was a time of important racial and
cultural change. During it, the Neanderthals were replaced by
Upper Paleolithic people similar to modern Europeans, and the
Mousterian flake culture was succeeded by a blade culture that
endured, in many forms and under many names, to the end of the
Pleistocene, around 8,000 b.c. Similar but not indentical blade
cultures have been found in Siberia, in northern Afghanistan, in
the Zagros Mountains of Iraq and Iran, in Turkey, and in Leba-
non, Syria, and Palestine.
A favorite cliche of anthropology, as widespread as the image
of the brutal Neanderthals, is that Upper Paleolithic Europeans
belonged to three races: the Cro-Magnon, which was Caucasoid;
the Negroid Grimaldis; and the Eskimoid race of Chancelade.
This concept is a product of the type-specimen procedure. There
was, in fact, only one Upper Paleolithic European race. It was
Caucasoid and it inhabits Europe today. We know this not only
from skeletons but also from the representations of the human
body in Upper Paleolithic art.
There was, as well, in the broad sense only one culture, al-
though archaeological splitters, after the fashion of their zoologi-
cal brethren, as defined in Chapter 1, are constantly dividing,
subdividing, and recombining it, treating the divisions as separate
cultures. Where sites are abundant, as in France, subdivisions of
cultures appear, vanish, and reappear in a manner perplexing
578
The Caucasoids
even to specialists. We are led to wonder whether these sequences
indicate invasions, diffusions of new techniques of making tools,
or a combination of both. I feel that the local populations re-
mained fairly constant but that genes flowed freely enough from
one region to another to prevent the rise of genetically different
races inside the Caucasoid subspecies. My concept of racial and
cultural homogeneity seems to be supported by the fact that Up-
per Paleolithic art styles show a remarkable continuity over a span
of 20,000 years.
In my opinion, the origins of the Upper Paleolithic culture have
been determined in a general way, but not all professional ar-
chaeologists agree with me. I believe that this culture and its ac-
companying racial type were imported into Europe and could
have come only from the East. Claims have been made that the
culture arose from a Mousterian prototype in Hungary, but the
only part of the Old World in which a blade culture is known to
have arisen from a flake culture in Wiirm I is the Near East —
Palestine, Syria, Lebanon, and possibly western Iran.9
Also, the makers of the blade tools were modern men, similar
enough anatomically to the Upper Paleolithic Europeans to have
been their ancestors. Palestine, Lebanon, and Syria are on the
Mediterranean coast, and anyone who walked west and then
north along the shores of Anatolia would soon find himself either
in Greece or on the shores of the Black Sea. This is the water-level
route that makers of hand axes followed in the Second Inter-
glacial; and makers of blade tools could just as well have traveled
it in the Gottweig Interstadial.
This is a logical and attractive theory, but it is too soon for us
to adopt it without reservation. In Turkey, Iran, and Afghanistan
many caves remain to be excavated, and who knows what will
turn up in them? It is not too soon, though, for us to feel that in
turning to the East we are on the right track.
9 For a detailed review of the extensive literature on this subject, including the
pioneer work of Dorothy Garrod, see:
ffowell: “Upper Pleistocene Stratigraphy and Early Man in the Levant,” PAPS,
Vol. 103, No. 1 ( 1959), pp. 1-65.
E. Anati: Palestine Before the Hebrews (New York: Alfred A. Knopf; 1962).
For the Iranian evidence, see also R. J. Braidwood, B. Howe, and C. A. Reed:
“The Iranian Prehistoric Project,” Science, Vol. 133, No. 3469 (1961), pp.
2008-10.
579
Upper Paleolithic Sites in Space and Time
According to Movius’s interpretation of about 120 Carbon- 14
dates,1 the Gottweig Interstadial began about 40,000 b.c. and
ended about 29,000 b.c. At its end intense cold set in. This marked
the beginning of Wiirm II, or, as Movius calls it, the Early Phase
of the Main Wiirm. It lasted about 2,000 years. Then came a long
interval of generally cold conditions, known as Wiirm III or the
Late Phase of the Main Wiirm. This lasted some 17,200 years and
was followed by two short cycles, each of which consisted of a
mild and a cold episode, totaling about two thousand years and
ending with the close of the Pleistocene, about 8,000 b.c.
According to Movius’s reconstruction, there were four main Up-
per Paleolithic industries in the Dordogne region, which is the key
area for Europe, because it contains the most complex sequence
and because more Upper Paleolithic digging has been done there
than anywhere else. First came the Perigordian, then the Auri-
gnacian, then a foretaste of the Magdalenian, then the final Auri-
gnacian, followed by the Solutrean, and then the rest of the
Magdalenian.
The Perigordian was a local industry, the oldest in that region.
The Aurignacian was a widespread industry, ranging from Spain
to Russia, and in central Europe it was probably as old as the Peri-
gordian was in France. The Solutrian was also widespread but
sporadic, with centers in Spain and Hungary, and the Magda-
lenian, which was probably derived from the Aurignacian, was
also widespread. In England there was only one industry, the
Creswellian, a local equivalent of the Aurignacian, and it lasted
until the end of the Pleistocene.
Upper Paleolithic Sites in Space and Time
Table 30 2 lists the Upper Paleolithic sites that have yielded
human skeletal material. I have not stated which bones were
1 Movius: Radiocarbon Dates and Upper Paleolithic Archaeology,” CA,
Vol. l , No. 5-6 (i960), pp. 355-91.
2 All but four sites in this list may be found in the Catalogue des Hommes
Fossiles of Vallois and Movius. The four exceptions may be found in:
(1) Movius and Vallois: “Crane Proto-Magdalenien et Venus du Perigordien
Final Trouves dans l’Abri Pataud, Les Eyzies (Dordogne),” L’Anth, Vol. 63, No.
3-4 (i959), PP- 213-32. A complete female skull, proto-Magdalenian.
(2) D. Ferembach: “Note sur une Mandibule Presumee du Magdalenien III,”
580 The Caucasoids
TABLE 30
UPPER PALEOLITHIC FOSSIL MAN SITES
COUNTRY CULTURE
COUNTRY CULTURE
Germany — 12
Limeuil, Dordogne M
Andernach am Rhein
M
Lussac-le-Chateau, Vienne M
Fiihlingen, near Koln
A
*La Madaleine, Dordogne M
Honert, nr. Dortmund
A
Le Ruth, Dordogne M
Kleine Scheuer, nr. Stuttgart
M
Massat, Aritige M
Neuessing, nr. Regensburg
S?
Montconfort, Haute-Garonne M
‘Oberkassel, nr. Bonn
M
Montesquieu- Avant6s, Ariege M
Petersfels, nr. Nordlingen
M
Pair-non-Pair, Gironde S
Ranis, nr. Weimar
M
*Le Roc, Charente S
Rothekopf, nr. Freiburg i. B.
M
La Rochette, Dordogne A
Sirgenstein, nr. Ulm
A
Les Rois, Vienne A
*Stettin, nr. Ulm
A
Roset, Tarn S
Ursprung, nr. Ulm
M
St.-Germaine-la Riviere,
Gironde M
Benelux — 6
St.-Vincent-Arlay, Rhone M
Hengelo, Netherlands
?
‘Solutrfj, Sa6ne-et-Loire A or S
Chaleux, Belgium
M
Ttioule, Haute-Garonne M
Goyet, Belgium
M
Terrasson, Dordogne M
Magrite, Belgium
A
‘Veyrier, Haute-Savoie M
Reviaux, Belgium
M
Oetrange, Luxemburg
A or M
Britain — 7
‘Aveline’s Hole, Somerset C
France — 43
Barcombe Mills, Sussex C
*Abri Pataud, Dordogne
M
Flint Jack’s Cave, Somerset C
Aurenson, Hautes-Pyren^es
M
*Gough’s Cave, Somerset C
Badegoule, Dordogne
SorM
‘Kent’s Cavern, Devon C
Blanchard, Dordogne
A
Paviland, Glamorgan C
Bourdeilles, Dordogne
S
Whaley, Derby C
Brassempouy, Landes
M
*Bruniquel, Tarn-et-Garonne
M
Spain — 7
‘Cap Blanc, Dordogne
M
Barranc Blanc, Valencia S
‘Chancelade, Dordogne
M
Carmago, Santander A
La Combe, Dordogne
A
Castillo, Santander A
‘Combe Capelle
A
Cobalejos, Santander M
Les Cottas, Vienne
A
Morin, Santander M
‘Cro-Magnon, Dordogne
A
‘Parpallo, Valencia S
Duruthy, Landes
M
Serinya, Gerona M
Entzheim, Haut-Rhin
?
Espalungue, Basses-Pyr6nees
M
Switzerland — 1
L’Espelungue, Landes
M
‘Bichon, Neufchatel M
Les Eyzies, Dordogne
M
Farincourt, Haute-Marne
M
Italy — 2
Grotte des F 6es, Gironde
M
* Arena Candide, Savona A
*Gourdan, Haute-Garonne
M
* Bdoussi R&oussi, Liguria A
*Les Hoteaux, Ain
M
Grotte des Enfants, Grimaldi
Isturitz, Basses Pyrenees
A or M
La Cave, Lot
S
Austria — 1
Laugerie-Basse, Dordogne
M
Miesslingtal, Lower Austria A
>> g | g W g g > 1** W OOOOOOO | g g g § g g Go;f*;»GeGoggggggg
58i
Upper Paleolithic Sites in Space and Time
TABLE 30 ( continued )
COUNTRY CULTURE
COUNTRY
CULTURE
Czechoslovakia — 8
Iran — 1
* Brno, Moravia
A
* Hotu Cave, Mazandaran
0
* Dolni Vestonice, S. Moravia
A
Dzerava Sk&la, W. Slovakia
S
Palestine — 5
* Mlade8, N. Moravia
A
Erg al-Ahmar
0
Podbaba, Prag, Bohemia
A
Jebel Qafza
0
* Predmosti, N.E. Moravia
A
Skhul
0
Sv. Prokop, Prag, Bohemia
A
Mugharet al-Wad
0
ZlatV Kuii, Central Bohemia
A
Mugharet al-Kebara
0
Hungary — 4
Barla-Barlang, Borsod, N.
Code to Cultural Symbols:
Hungary
M
A = Aurignacian,
incl.
Perigordian
Czakvari-Barland, Fejer, W.
S = Solutrian
Hungary
M
M = Magdalenian
* Nagy-Sap, Esztergom, N.W.
C = Creswellian
Hungary
M
0 = Others
Pilisszantou-Kofulke, near
Budapest
M
Complete Skulls or Skeletons Described
Numbers of Sites
Rumania — 1
by Countries
by Cultures
Cioclovina, Transylvanian
Germany
2
A = 9
Alps
A
France
11
S = 3
Britain
3
M = 10
U.S.S.R.— 6
Spain
1
C = 3
Puskari, Ukraine
0
Switzerland
1
0 = 1
Tsulatovo, Ukraine
0
Italy
2
26
Korman, N. Bessarabia
A
Czechoslovakia
4
Siuren’i, Crimea
A
Hungary
1
Devis-Khreli, Georgia
A
Iran
1
Mal’ta, Siberia
0
26
* Published studies of complete skulls or skeletons are available.
found in each site because I do not intend to go over all the ma-
terial in detail. The sites for which published studies of complete
skulls or skeletons are available have been starred.
The geographical distribution of these sites resembles that of
BSA, Vol. 5 ( 1954), pp. 25-34. A mandible from St.-Vincent at Arlay, Lyon; Late
Magdalenian.
(3) M-R. Sauter: “Etude des Vestiges Osseux Humains des Grottes Prehis-
toriques de Farincourt (Haute-Mame, France),” ASAG, Vol. 22, No. 1 (1957),
pp. 6-37. A late Magdalenian mandible and maxilla from Farincourt.
(4) Sauter: “La Squelette Prehistorique de la Grotte du Bichon (Cotes-du-
Doubs, La Chaux-de-Fonds, Neuchatel),” AS, Vol. 9, No. 3 (1956), pp. 330-5.
A Magdalenian skeleton from the cave of Bichon, Neufchatfll, Switzerland.
582
The Caucasoids
the Neanderthals. Places too cold for comfort in Wiirm I were also
difficult to live in during Wiirm II, III, and later. Southern France
was again the favored spot. In Germany only the Rhineland and
western Bavaria were popular, and most of the Czechoslovakian
skulls are from the Interstadial.
Of 161 listed sites, only the remains of 25 have been adequately
described in publication. Eleven are from France, and, by coinci-
dence, eleven of these skulls or skeletons are Magdalenian. Not
one was found east of Moravia and northwestern Hungary. Many
of the publications are so old that the measurements were not
based on standard techniques; and old calculations of stature are
usually much too high. Two physical anthropologists, G. W.
Morant and G. von Bonin, have remeasured and reworked as
many skulls and long bones as possible; their monographs princi-
pally document the following survey.3
The Racial Characteristics of the
Upper Paleolithic Europeans
The Upper Paleolithic Europeans, who lived from about 30,000
to about 10,000 years ago, were modern Caucasoids. Were they
barbered and dressed in the current styles, they could sit in any
western European restaurant without arousing particular com-
ment except for their table manners. A few very observant fellow
customers might notice that they closed their deeply worn teeth
with an edge-to-edge bite, and that their well-developed temporal
and masseter muscles bulged as they chewed.
As von Bonin has shown, the men were not notably tall. The
mean stature for twelve adult male skeletons is only five feet eight
inches (173 cm.), shorter than modern Americans. The famous
Old Man of Cro-Magnon, depicted in textbooks as a giant, was
only five feet six ( 168.4 cm. ) • The two tallest men of the series,
Grotte des Enfants and Barma Grande 2, were five feet eleven and
a half inches ( 181.8 cm). The shortest Upper Paleolithic man was
3 Morant: “Studies of Palaeolithic Man, IV, A Biometric Study of the Upper
Palaeolithic Skulls of Europe,” AE, Vol. 4 (1930), pp. 109-214.
G. von Bonin: “European Races of the Upper Paleolithic,” HB, Vol. 7, No. 2
(i935), PP- 196-221.
Racial Characteristics of the Upper Paleolithic Europeans 583
Chancelade. He was only five feet three ( 160 cm.),4 and he lived
during a brief spell of intense cold. As we know, intense cold tends
to reduce stature.
The five female skeletons have a mean stature of five feet one
inch (155 cm.), and the range is only from 154 to 157.5 cm.
Females were, then, much shorter than the men. The sex differ-
ence was probably real, and not merely an accident of sampling in
a small series, because the women’s skulls are also much smaller
than the men’s.
The long bones of these skeletons are on the whole slender, like
those of Krapina and Mount Carmel, and a lean body build is
indicated. In two respects the long bones of these skeletons are
variable — in the proportion of the length of the forearm to the
length of the humerus, and in the ratio of the thigh to the lower
leg. The Aurignacian Grimaldi woman, found in a double burial
with her so-called Negroid son, had a long radius and a short
humerus. Elongated shin bones were found in the skeleton of
Combe Capelle, who lived in the mild Gottweig Interstadial, and
in two Aurignacian skeletons from the Riviera, one from Grotte
du Cavillon and the other ( not to be confused with the Grimaldi
pair) from Baoussi Raoussi.5 None of these men, nor the Grimaldi
mother and child, was exposed to great cold. Both Skhul 4 and
Skhul 5 had similar limb proportions.
The hands and feet of the Upper Paleolithic Europeans are
better known to us from archaeological than from osteological evi-
dence. Many negative silhouettes of hands, made by spraying pig-
ment out of a bone tube over a hand held against a wall, have
been found on the walls of French caves; bare footprints have been
found on cavern floors in France and Italy. Both the hands and
the feet were normal for slenderly built Europeans.
Morant’s series of twenty male skulls and von Bonin’s series of
thirteen female skulls represent nearly all countries from France
4Vallois: “Nouvelles Recherches sur la Squelette de Chancelade,” L’Anth,
Vol. 50, No. 1-2 (1941-6), pp. 11-202. Vallois calculated stature from the hu-
merus, femur, and tibia; von Bonin, who got a lower figure, used the humerus
only.
5 Von Bonin also found high humeroradial and femorotibial indices for the
Magdalenian skeleton of Obercassel, but he doubted the accuracy of the original
measurements.
The Caucasoids
584
to Czechoslovakia, and all archaeological cultures. The male
skulls are large, with a mean cranial capacity of 1,580 cc.; the
female skulls are much smaller, with a capacity of 1,370 cc., about
the size of Swanscombe. The detailed measurements of these
skulls reveal a very long, moderately broad and high brain case
of fully modern proportions; a face of moderate to great length,
and a considerable breadth. In fact, the bizygomatic diameter ex-
ceeds the cranial breadth. This was not the case among the Ne-
anderthals, nor is it among most modern Europeans. Heavy chew-
ing, combined with a relatively narrow brain case, is responsible
for this archaic feature, found also among the Eskimo. It has no
racial significance.
Most of the skulls are not prognathous, an exception being the
recently discovered female skull from the Proto-Magdalenian of
Abri Pataud. Brow ridges are of moderate size or they are missing
in most of the skulls, except in the Czechoslovakian ones of the
early Aurignacian, which come closest to Skhul 5 and Jebel
Qafza 6. They, however, show no trace of Skhul 5’s alveolar prog-
nathism. The series of male skulls also resembles a later, Neolithic
series from France, Iron Age skulls from Norway, and Anglo-
Saxon ones from the east coast of England.
Although most of the teeth are too worn to permit accurate ob-
servation, those of the so-called Negroid boy of Grimaldi are in
perfect condition. His upper jaw shows irregular tooth eruption,
gaps, and malocclusion. The upper median incisors have vertical
ridges on the lingual side, and a basal protuberance. These are
dental characteristics of the Negro, but not exclusively. They are
also seen on a number of teeth from Krapina and on those of Ne-
anderthals, and are also present, as we have just mentioned, in the
Mount Carmel population. An upper canine from the Magdalen-
ian maxilla of Farincourt6 has the same features. The Grimaldi
child was no more Negroid than the Palestinians of Skhul and
many living Europeans of the Mediterranean region.
The other alleged intruder in the European population, Chan-
celade, had wide zygomatic arches and flaring gonial angles, as
befitted a heavy chewer living in extreme arctic conditions. But he
6 Sauter: “Etudes des Vestiges. . . .”
Racial Characteristics of the Upper Paleolithic Europeans 585
had high-rooted, aquiline nasal bones; a face that was far from
flat; and a completely Caucasoid configuration of the malars.7 He
was as European as the rest of the Upper Paleolithic people.8
The Upper Paleolithic Europeans were great artists: they
worked in bone, ivory, antler, and limestone and carved in the
round and in relief, and engraved and painted. But they were
interested more in depicting animals than in depicting people.
Very few human faces and figures appear in any of the media. Of
these, some are exaggerated, others are humorous, and a few are
realistic.9 The statues in the round and the bas-reliefs, known as
“Venuses,” invariably represent grossly obese women, whose fat
is deposited on the same parts of the body and in the same fashion
as in living fat women of European origin, many of whom also
have slender bones.
A unique wall engraving in La Magdaleine cave shows a long-
breasted woman with tapering extremities, her hips and waistline
only a little fuller than is currently fashionable. Another notable
wall engraving, found in Sicily, depicts some kind of ceremony;
the men taking part have normally proportioned Caucasoid
bodies.
Wall paintings, wall engravings, and ivory carvings contain a
number of portraits of human faces. Some are bearded, some bald.
Most of the men shown have prominent noses. One woman has
7 Morant: “Studies of Palaeolithic Man, I, The Chancelade Skull and its Rela-
tion to the Modem Eskimo Skull,” AE, Vol. l (1926), pp. 257-76.
Vallois: “Nouvelles Recherches. . . .”
8 There is, however, a possibility that a few North Africans may have visited
Europe during Wiirm II or III. This is indicated not only by the discovery of
Aterian arrowheads in Solutrean deposits in Spanish caves, mentioned on page
523, but also by the discovery of a skullcap, cut in the form of a bowl, which was
found lying on the floor of a cave containing Upper Paleolithic implements and
paintings. It has very heavy brow ridges and a receding forehead, and could
hardly have belonged to an Upper Paleolithic Caucasoid. In the only available
photograph it looks, in profile, like the Florisbad skull from South Africa, an
ancestral Bushman specimen of a group which probably originated in North
Africa. Until this skullcap has been studied, no definite statement about it can
be made. A. H. Brodrick: “A Newly Discovered and as yet Unexplored Treasure-
House of Spanish Cave Art: The Fantastic and Beautiful Caves of Nerja — a
Preliminary Note.” ILN, Vol. 239, No. 6366 ( 1961), pp. 216-9.
9 Paolo Grazioso: Palaeolithic Art (New York: McGraw-Hill Book Co.; i960).
J. A. Mauduit: 40,000 An. s d’ Art Moderne (Paris: Librairie Plon; 1954).
586
The Caucasoids
Fig. 76 The Human Face and Hand in Upper Paleolithic Art. These en-
gravings made on the walls of French caves are probably all of Magdalenian, or
Late Upper Paleolithic, origin. They have been selected from a large number of
copies, some of which are of doubtful authenticity. They show both that Upper
Paleolithic Europeans were Caucasoid and that they had a sense of humor. ( Draw-
ings after P. Graziosi, i960. )
nasal prognathism to a marked degree, and another is chinless.
Some of these drawings reflect to a certain degree the imagination
of modem archaeologists who copied them by lamplight in the
depths of the caves; but most are accurate. The total effect is that
of a set of caricatures of modern Europeans.
T heir Asiatic Relatives
587
The Fate of the Upper Paleolithic Europeans 9
The Upper Paleolithic Europeans did not vanish with the mam-
moths on whose succulent flesh they feasted, nor with the Ne-
anderthals. They survived the Pleistocene, and their descendants
became Mesolithic salmon-seiners, Neolithic villagers, Bronze Age
warriors, and Iron Age Vikings. They followed the reindeer to the
edge of the ice, and when it melted, there they remained. But they
were a restless people, and their descendants still are. After they
had learned agriculture and cattle breeding from others like them
who had come from the East, they expanded, migrating south-
ward and eastward in many waves, one of which even reached
India; and their descendants are to be seen in America, Australia,
New Zealand, and South Africa.
Their Asiatic Relatives
Very little skeletal material is available from the Upper
Paleolithic sites in western Asia, largely because it has not been
looked for. However, from the cave of Hotu, on the Caspian shore
of Iran in west-central Asia, we do have three skeletons that date
from the penultimate millennium of the Pleistocene.1 In brief, this
man and two women from Hotu were indistinguishable from their
western European contemporaries.
What little we have from Palestine, mostly scraps of bone and a
few teeth, is also Caucasoid. For example, the Mesolithic Natufian
skulls and long bones from that country are those of ancestral
Mediterraneans.2 As we shall see in the next chapter, some of the
Near Eastern Caucasoids invaded North Africa before the Pleis-
tocene was over. Others, remaining in western Asia, were the first
people to grow crops and to tame the ancestors of our domestic
animals. The Neolithic culture that they had invented spread in
many directions, and became the basis of our modern civilization.
9 If I may do so without immodesty, I recommend my Races of Europe for a
review of this subject.
1J. L. Angel: “The Human Skeletal Remains from Hotu Cave, Iran,” PAPS,
Vol. 96, No. 3 ( 1952), pp. 258-69.
2 McCown: Natufian Crania from Mt. Carmel (Berkeley, California: Univer-
sity of California Library; 1940).
AFRICA
S.
The Darkest Continent
Several years ago a number of old friends and neigh-
bors sat in my house talking. Among them was Sarah Jones, a
Negress born shortly after emancipation. Two of the group, being
New Englanders, were, as might be expected, discussing ances-
tors. Mrs. Jones listened intently. Then she turned to me and
asked: “Professor, who were my ancestors?”
I had to reply that I did not know. Thanks to Lewis Leakey and
his recent discoveries, I know a little more now than I did then,
but not enough to be certain. The origin of the African Negroes,
and of the Pygmies, is the greatest unsolved mystery in the field of
racial study. In this chapter I shall present all the evidence I can
find and offer a tentative solution.
To begin with, Africa is not the home of one subspecies but of
two. The Bushmen evolved there as well as the Congoids — Ne-
groes and Pygmies. The Caucasoids of North Africa, Berbers and
Arabs, are late arrivals. When the southern Mongoloids were in-
vading southeast Asia and Indonesia, the ancestors of the Berbers
invaded North Africa, pushing the earlier inhabitants southward,
just as the Australoids were crowded eastward and southward
over Wallace’s Line into their present home.
In Asia and Indonesia the question of who crowded whom is
easy to answer because there was only one invader and only one
displaced group. In Europe the same was also true of the Neander-
thals and of the Upper Paleolithic peoples who replaced them.
But in Africa two subspecies were displaced by the Caucasoid in-
vaders. Our first problem is to discover where each of the native
The Darkest Continent
589
races lived before the invasions, and where each went afterwards.
In recent centuries Negroes have inhabited most of Africa be-
tween the Sahara and the Limpopo River, whereas Bushmen and
their cattle-breeding kin, the Hottentots, have occupied South Af-
rica and parts of southern Rhodesia. The boundary between Ne-
groes and Bushmen is not an impenetrable geographical barrier
but a clinal region, and the two subspecies could not have evolved
each on its own side of it because the isolation needed for sub-
specific evolution did not exist. By simple zoogeographical logic
we must therefore assume that at least one of the two subspecies
initially moved into its present territory after each had evolved,
during the Pleistocene, in a state of comparative isolation. And it
is easier to believe that one moved first than that both moved
simultaneously. Obviously, the subspecies that felt the Caucasoid
pressure first moved first and farthest.
The Pygmies hold the key to the problem because North Af-
rica is not the kind of country in which Pygmies could have
evolved. It is not, and never has been, a tropical forest region. The
present home of the Pygmies is, quite appropriately, the rain forest
of the Congo and of sections of West Africa, and it is the logical
place for them to have evolved in. During parts of the Pleistocene
the Congo Basin was under water, and the ancestors of the Pyg-
mies must then have lived in the edges of the forest, and have
entered the more elevated parts of that refuge in times of drought.
Once in the forest, they became dwarfs, according to the rules
governing dwarfing as discussed in Chapter 3.
Pygmies are obviously related to Negroes, and a full-sized
Pygmy ancestor and a Negro ancestor of the same period may
have been indistinguishable. The Negro homeland must therefore
have been the savannahs at the edge of the forest, an environment
to which Negroes are physiologically adapted. They could not
have acquired their ability to withstand heat, particularly damp
heat, during the 12,000 years since the Caucasoids pushed their
predecessors out of North Africa. The South American Indians of
the Amazon basin have failed to become heat-adapted in an equal
length of time. The African forest and its peripheries are there-
fore the Congoid home.
The Bushmen, who are not heat-adapted, do not fit this picture.
590
Africa
But, as we shall presently see, there is fair evidence that the ances-
tors of the Bushmen were full-sized people and that they evolved
in North Africa, north of the Saharan barrier which gave them the
isolation they needed to become a separate subspecies. When the
present Palearctic fauna invaded North Africa near the end of
the Pleistocene, the Caucasoids who came with it drove out the
Capoids, who crossed the Sahara via the central Saharan Tibesti
highlands, and then followed the cool East African highlands
southward to their present home.
There they entered an underpopulated area inhabited by hu-
man beings of a lower evolutionary grade, who were related to the
ancestors of the Negroes and Pygmies living farther north and
west. These aborigines gave the ancestors of the Bushmen little
trouble, and were absorbed by the invaders. Much later, in full
historical times, some of the Negroes of West Africa who had
acquired agriculture and iron metallurgy moved eastward and
southward and in turn absorbed many of the Bushman tribes.
They arrived in South Africa simultaneously with the Dutch.
Bantu and Boer then formed the jaws of a giant pincers that drove
the Bushmen into the Kalahari and led to the racial conflicts that
beset that troubled land today.
This is the most plausible outline of African racial history that
it seems to me can be drawn from available evidence. In the rest
of this chapter I shall try to document this outline.
Fossil Man in North Africa: the Ternefine-Tangier Line
As stated in Chapter 7, Lower Pleistocene archaeological
sites are as old in North as in East Africa. Moreover, what may be
the oldest Australopithecine yet found comes from the heart of
the Sahara, in the Bepublic of Tchad, halfway between these two
most ancient archaeological regions. Thus, North Africa has as
good a claim to the title of Cradle of Mankind as Tanganyika.
It is therefore disappointing that we have no North African
Australopithecine or human remains older than the Early Middle
Pleistocene and that for the vast period between then and the ar-
rival of the Mouillians, shortly before the end of the Pleistocene,
The T ernefine Discoveries 591
TABLE 31
PRE-MOUILLI AN SKELETAL MATERIAL
FROM NORTH AFRICA
Country
Site
Age
Material
Name
Algeria
Temefine, Oran
Early Middle
Pleistocene
1 piece parietal
3 mandibles
Atlanthropus
mauritanicus
Morocco
Sidi Abd er-Rahman
Riss or Third
1 piece mandible
none
Casablanca (Lito-
rina Cave)
Pluvial
Temara (Smugglers’
Early Last
1 piece mandible
none
Cave)
Interglacial
Rabat
Late Last
1 piece mandible
none
Interglacial
1 piece maxilla, frag-
ments cranial vault
Mugharet al-‘Aliya
End Last
1 fragment child’s
none
Tangier (High Cave)
Interglacial to
Wiirm I
maxilla
Taforalt
Prob. Wiirm I
1 piece calva
none
Libya
Haua Fteah
34,000 BP
(Gottweig
1 fragment mandible
H. neander-
Interstadial)
thalensis
all we have is seven mandibles, some quite fragmentary, a piece
of an adult maxilla, another from a child, and several small pieces
of cranial vault ( see Table 3 1 ) .
The T ernefine Discoveries 1
I n 1954 and 1955 Camille Arambourg, a renowned French
paleontologist, discovered human remains in a rich deposit of fos-
1 C. Arambourg: “L’Hominien Fossile de Temefine (Algerie),” CRAS, Vol.
139 (1954), PP- 893, 895.
Arambourg: “A Recent Discovery in Human Paleontology, Atlanthropus of
Temefine (Algeria),” AJPA, Vol. 13, No. 2 (1955), pp. 191-6.
Arambourg: “Une Nouvelle Mandibule ‘d’Antlanthropus’ du Gisement de
Temefine,” CRAS, Vol. 241 (1955), pp. 431-3.
Arambourg: “Le Parietal de l’Atlanthropus Mauritanicus,” CRAS, Vol. 241
(1955), PP- 980-2.
Arambourg: “Une Illme Mandibule ‘d’Atlanthropus’ Decouverte a Temefine,”
Quaternaria, Vol. 3 (1956), pp. 1-4.
Arambourg: “Recentes decouvertes de paleontologie humaine en Afrique du
Nord frangaise,” PTPA, 1957, pp. 186-94.
F. C. Howell: “European and N. W. African Middle Pleistocene Hominids,”
CA, Vol. 1, No. 3 (i960), pp. 195-232.
592
Africa
sil animal bones in a flooded sandpit at Ternefine near Palikao,
eleven miles ( 17 km. ) southeast of Mascara in the Department of
Oran, Algeria. The Early Middle Pleistocene date was deter-
mined by examination of the fauna, which was typically African
and indicated a savannah type of landscape. The associated in-
dustry was early Acheulian mixed with many choppers, chopping
tools, and flakes reminiscent of the Far East. It seems to have been
a generalized early industry into which the making of hand axes
Fig. 77 The Ternefine Parietal. The only piece of skull recovered at Ternefine
was this parietal, shown from the inside. No scale was given, but this drawing is
exactly the same size as that published. The middle meningeal artery pattern
suggests that it belonged to a member of Homo erectus. ( Drawing after Arambourg
1955- )
had been introduced, or in which hand-axe manufacture had
been invented. The human remains consisted of a right parietal
bone and three mandibles.
The right parietal bone belonged to an immature individual.
We know this because all the sutures are open, and the bone is no
thicker than that of a modern adult. Its curvature suggests a
low vault, with the maximum cranial breadth lying below its junc-
ture with the temporal. Although the dimensions of this bone have
not been published and the drawing 2 has no scale, I have been
2 Arambourg: “Le Parietal. . . .”
The T ernefine Discoveries
593
told by two professionals who have handled it that it lies some-
where between Sinanthropus and Neanderthal in size, and that it
approximates the size of Swanscombe’s right parietal.
The size as stated does not tell us whether the bone belonged to
a large Homo erectus skull or a small Homo sapiens one, but the
morphology and the endocranial surface configuration suggest
the former. Like the Pithecanthropi and Sinanthropi, it has a
prominent Sylvian crest and a simple meningeal artery pattern.
Mandible 1 is nearly complete, with all its molars and pre-
molars and its right lateral incisor. Number 2 consists of an en-
tire left side and enough of the right side to allow room for the
two right incisors, but its only teeth are its left molars and pre-
molars. Number 3 is complete except for the post-mortem loss of
seven teeth; those present are the right lateral incisor, right ca-
nine, left first premolar, both second premolars, and all six molars.
Mandible 3, which Arambourg classifies as masculine, is the
largest lower jaw yet found which all investigators agree is hu-
man. Its bicondylar breadth is very great, for example, so great
that the cranium which it fitted must have had a very wide base —
an erectus feature.
In many respects Number 3 is almost Australopithecine. In fact,
it resembles the Swartkrans mandibles in its dimensions and also
in one particular morphological detail. Its ascending ramus is
very high and inclined far backward, with a 70 0 angle of inclina-
tion; and the coracoid process of its ascending ramus is higher
than its condyle. Temefine 3 shares this overall configuration of
the ascending ramus, to a lesser degree, with its contemporary,
Sinanthropus, and the Neanderthals, who lived much later. The
low broad ascending ramus of the Heidelberg jaw is so different
that it must represent an entirely separate evolutionary line.
The other two Ternefine mandibles are similar to Number 3 in
shape but they are much smaller, so much so that a sexual dimor-
phism is suggested for the North African population, as among the
Sinanthropi. Mandible 1 has two mental foramina; Number 3
has two on the right and three on the left; and the one-sided
Number 2 has a single foramen. In these respects the Ternefine
jaws resemble both Sinanthropus and the Neanderthals.
Despite its size, Ternefine 3 does not have the largest teeth;
594
Africa
these are found in the jaw of Number 2, supposedly a female.
Her molars and premolars are larger than those of Sinanthropus,
and the first and second molars are larger than those of Pithecan-
thropus B, who lived more than 100,000 years earlier. The molars
and premolars of Ternefine 1 and 3 fit comfortably within the Si-
nanthropus range and are a little smaller than the Pithecanthro-
pus B teeth. However, in the Pithecanthropus B mandible the
third molar is the largest, followed in order by the second and
then the first molars, whereas in all three Ternefine jaws the sec-
ond molar is the largest and the third the smallest, as in Sinan-
thropus.
In all three Ternefine jaws the incisors and canines seem small
in relation to the premolars and molars, but this is difficult to
establish because these teeth are either badly worn or absent al-
together and represented only by sockets.
Fig. 78 Mandibles: Tebnefine 1 and Rabat. The Ternefine mandibles are not
all alike. No. 1 (A) has a steep, rounded sagittal profile. The Rabat fragment
( B ), probably 200,000 years younger, is still steep, but has the beginnings of a chin.
The premolars and molars are heavily wrinkled, as in Sinan-
thropus and some of the Australopithecines, and they are tauro-
dont. Nearly all have basal cingulums, and all the molars have the
Y-5 or +5 cusp pattern. The canine of Number 3 has a long, thick
root. The form of the incisors cannot be determined. All in all,
these teeth resemble those of the Australopithecines, Pithecan-
thropus, and Sinanthropus, but the closest resemblance is to Si-
nanthropus. Except for taurodontism, they have little in common
with the Heidelberg teeth.
The Ternefine specimens are important and tantalizing. The
The Mandible from Smugglers’ Cave, Temara, Morocco 595
skull was apparently erectus in general form, but large enough to
have been either erectus or sapiens ; the face, judging by the
length of the ascending rami of the mandibles, was very long, and
the jaws themselves formed a bridge between those of the larger
Australopithecines and the Homines erecti of China and Indo-
nesia.
The Litorina Cave Mandible
I N 1953 a French archaeologist, P. Biberson, found human re-
mains in a former cave in the quarry of Sidi Abd er-Rahman
(named for a saint’s tomb perched atop it) in the Anfa section of
Casablanca, Morocco. The culture was an evolved Acheulian in-
dustry, and the deposit that of the so-called Tyrrhenian I period,
identified by means of associated sea-levels, which fluctuated on
the Moroccan coast during the Pleistocene. It probably coincided
with the Riss glaciation in Europe.
The specimens consist of two small pieces of mandible contain-
ing three right molars and a left first premolar.3 Morphologically
these fragments resemble those of Ternefine, but they are a little
smaller. Had we three Litorina Cave jaws to match the three of
Ternefine, we might find no difference at all.
The first premolar has a cingulum, and the molars are wrinkled.
The first two molars have five cusps each, and the third one six.
The size gradation of the three molars is second, first, and then
third, as in Ternefine and Sinanthropus.
Despite the time gap of about 200,000 years, the genetic con-
tinuity between Ternefine man and that of the Litorina Cave
seems just as clear as their cultural continuity.
The Mandible from Smugglers Cave, Temara, Morocco 4
During or before 1958, Father Jean Roche excavated a cave
called Grotte des Contrabandiers on the Moroccan coast at Te-
3 Arambourg: “Recentes decouvertes. . . .”
Howell: “European and N. W. African Pleistocene Hominids.”
V. Vallois and J. Roche: La Mandibule Acheuleene de Temara, Maroc ”
CRAS, Vol. 246 ( 1958), pp. 3113-6.
Africa
596
mara, 33 miles (53 km.) northeast of Casablanca. Among other
undescribed human remains, he found a mandible, nearly com-
plete except that parts of both ascending rami had been broken
off. The artifacts belonged to the final Acheulian industry or to
the threshold between the Acheulian and the succeeding flake
culture, the Aterian, and were roughly contemporaneous with the
Litorina Cave mandible.
The Smugglers’ Cave mandible resembles those of Ternefine
and the Litorina Cave fragments in most respects, but it is the
smallest yet found of the North African group. Unlike the others,
it had a nearly straight profile and as much chin as some of the
Neanderthals. The teeth are as large as those of the Litorina Cave
jaw. The first molar is the largest of the three, followed in turn by
the second and third.
The canine, which is very large, is ribbed on the lingual side
into a three-chambered surface, and its cutting edge is horizontal
rather than pointed. As among the Australopithecines, the second
premolar is molarlike in structure. All the molars are moderately
taurodont, and all of them have a Y-5 cusp pattern, except for the
right first molar, the pattern of which is +4.
The Rabat Remains 5
I N 19.33 quarrymen blasted what was probably a complete skull
from a sandy marine consolidation on the outskirts of Rabat. All
that was recovered, however, were portions of the lower and up-
per jaws and brain case, as follows: (1) the front half of a mandi-
ble containing three incisors, one canine, three premolars, and a
row of three molars; (2) the lower part of the right maxilla with a
small piece of palate, to which is attached a natural cast of most
5Vallois: “L’Homme Fossile de Rabat,” CRAS, Vol. 221 (1945), pp. 669-71.
M. Boule and Vallois: Les Hommes Fossiles (Paris: Masson et Cie; 1952),
pp. 443-4-
Vallois and Roche: “Le Mandibule Acheuleene. . . .”
Vallois: “L’Homme de Rabat,” BAM, Vol. 3 (i960), pp. 87-91.
L. C. Briggs: “The Stone Age Races of Northwest Africa,” BASF, Vol. 18
(i955), PP- 17-19-
Howell: “European and N.W. African Pleistocene Hominids.”
Bruce Howe, personal communication regarding date.
The Rabat Remains
597
of the rest of the palate, and two incisors, one canine, two pre-
molars, and two molars; and (3) twenty-one small fragments of
the cranial vault, not one of which is larger than a twenty-five-
cent piece. The date of this find is a period called Tyrrhenian I— II,
in shore-line chronology probably equivalent to the end of the
Last Interglacial in Europe and almost certainly no older than the
onset of Wiirm I.
The cranial fragments have not been, and probably could not
be, reassembled. But they are not much thicker than the mean for
modern skulls. On the basis of the sutures and of the teeth, the
skull is attributed to a seventeen-year-old male.
The maxilla lacks a canine fossa, and indicates pronounced
alveolar prognathism; the palate was large. The mandible, about
the size of the Temara specimen, is smaller than any of the three
Ternefine jaws, and morphologically is similar to the other North
African jaws in this series. It has two mental foramina on the right
side, and a large one on the left. Its symphyseal profile resembles
those of Ternefine, and its angle of inclination, 65°, is the same as
that of Ternefine 2.
The lower molars also fit the Ternefine range. The first is the
largest, followed by the second and the third. In size and shape
the molars resemble those from the other early North African sites,
except that the third molar has six cusps.
The upper incisors (the first we have seen from North Africa)
are shovel-shaped, although not to the degree found in Sinanthro-
pus, and the lower incisors form a nearly straight line from canine
to canine. The upper canines, like those of Sinanthropus, have
heavy cingulums on the outer sides of the base of each crown, and
the lingual surface is divided vertically by a double ridge. These
teeth are pointed and extend a little beyond the level of the in-
cisors. The lower canines, however, are incisorlike, as in the Ne-
anderthals.
The upper premolars have a complicated cusp pattern, as in
Sinanthropus; the first has two roots, the second a single long
root. The lower premolars are asymmetrical, like those of Swart-
krans, with diamond-shaped crowns. The first lower premolar has
a high lingual cusp, the second lower premolar has two roots and
a large distal portion; it is thus ‘ molarized” as in the Australo-
Africa
598
pithecines and Sinanthropus. Some of the Neanderthals share
these special dental features with Rabat man and Sinanthropus,
but to a lesser degree.
Tangier Man
Our last find in this series, dated at the base of Wiirm I, comes
from the northern end of the Moroccan coast and is about as old
as the Rabat specimen, or perhaps a little younger. I found it in
1.939 while excavating the High Cave (Mugharet al-‘Aliya), one
of the Caves of Hercules facing the Atlantic on Ras Ashagar, a few
miles south of Cape Spartel, to the southwest of Tangier. It was in
a layer of yellow soil underlying two other Pleistocene strata. All
three contained Aterian implements, but the refined bifacial
points equipped with tangs suitable for hafting as arrowheads,
and typical of the later stages of that culture, lay only in the layers
above my specimen.0
It was a piece of a child’s maxilla, with erupted and unerupted
teeth. Of these, a permanent canine and first premolar have been
measured. The child had died at about the age of nine. In sifting
the earth from the same layer, I also found a badly worn upper
first molar of an adult.
The piece of maxilla extends from the socket of the left first
permanent upper incisor to that of the unerupted second molar,
and includes the floor of the nasal aperture, the base of the nasal
wall, and a small piece of the zygomatic process. The bone is
massive, indicating a face already large and long at an early age;
the canine fossa is absent, and the lower border of the nasal mar-
gin is smoothly rounded, as in Negroes and Australoids. It re-
sembles in essential details the maxilla of the seventeen-year-old
from Rabat.
The upper canine and first premolar are large, large enough to
match those of Ternefine and the other early North Africans.
Whereas the canine is particularly thick labiolingually, the first
6 B. Howe and H. L. Movius, Jr.: “A Stone Age Cave Site in Tangier,” PMP,
Vol. 28, No. 1 (1947).
H. Hencken: “The Prehistoric Archaeology of the Tangier Zone, Morocco,”
PAPS, Vol. 92, No. 4 ( 1948), pp. 282-8.
M. S. $enyiirek: “Fossil Man in Tangier,” PMP, Vol. 16, No. 3 ( 1940).
Tangier Man
599
Fig. 79 The Tangier Maxilla and Teeth. B. Buccal view; D. Distal view;
L. Lingual view; M. Mesial view; O. Occlusal view. a. Lateral view of the Tangier
maxilla after the extraction of the teeth; b. Anterior view of the Tangier maxilla
after the extraction of the teeth; c. Left permanent upper canine of Tangier man;
d. Left permanent upper first premolar of Tangier man; e. Left upper second molar
of the Tangier man. Approximately natural size. Note that the maxilla is puffy, as
in Mongoloids and Neanderthals, and that there is no canine notch. The teeth, with
low crowns and stout roots, resemble those of Sinanthropus. (Drawings from
Senyiirek, 1940.)
premolar is relatively narrow in that dimension. The canine, like
that of the Rabat specimen, has a cingulum. It lacks the fingering
ridges of the labial side, which is smooth, but it has a marked
triangular eminence, or heel, found also in some Neanderthals.
The first upper molar of the second individual is so badly worn
that its cusp pattern cannot be detected, and its dimensions may
6oo
Africa
have been reduced by attrition. Still, it is a very large tooth, well
above the Sinanthropus mean and it probably was above the Si-
nanthropus maximum before wearing down. Indeed, it may well
have approached the dimensions of Pithecanthropus 4’s first
upper molar.
The Taforalt Cranial Fragment
The only other specimen of Aterian man yet found is a very
small piece of cranial vault found in an Aterian cultural level in a
cave at Taforalt, in the Beni Znassen country of northeastern
Morocco not far from Oujda.' It has not been described, and prob-
ably does not warrant description.8
The T ernefine-T angier Line, Cannibals, and Bushmen
A l l in all, the Tangier child and his older companion were true
successors of other North Africans from Ternefine, Litorina Cave,
and Smugglers’ Cave, and similar to their contemporary from
Rabat. Together these specimens form a single line. They were
certainly not Caucasoid, nor especially Negroid. They bear a sim-
ilarity on the one hand to the Australopithecines and on the other
to both Pithecanthropus and Sinanthropus, more particularly to
the latter.
They probably belonged to the erectus grade when they first
appeared, but whether they had achieved the sapiens grade by
the time the Caucasoid Mouillians invaded shortly before the end
of the Pleistocene is unknown. However, the implements from
later phases of the Aterian culture were sophisticated flake tools,
pressure-flaked on both sides, and some of them had tangs for
hafting. They were as good as the recent work of Bushmen, and
this circumstantial evidence suggests that the men who made
these tools were Homines sapientes.
The relationship of the Tangier child and his companion to
7J. Roche: “La Grotte de Taforalt,” L’Anth, Vol. 57, No. 3-4 (1953), pp.
375-8o.
8 Briggs: “The Stone Age Races. . . .”
jl
The T ernefine-T angier Line, Cannibals, and Bushmen 601
Pithecanthropus and Sinanthropus can be explained only on a
theoretical basis, since we do not know the antecendents of the
three populations. Either the Ternefine-Tangier people were de-
scended from immigrants from east Asia; or the ancestors of Pithe-
canthropus and Sinanthropus came from North Africa; or, as a
third possibility, the ancestors of all three fossil subspecies came
from some point geographically in between.
In view of what we know of Lower Pleistocene archaeology, the
most likely possibility is that all three originated in North Africa
and at an earlier evolutionary level than any yet seen in the skulls
of the genus Homo, but this theory cannot be proved or disproved
until more, and earlier, skeletal material is unearthed both in
North Africa and in Asia.
On the other hand, it now seems fairly likely that the Ternefine-
Tangier people had something to do with the origin of the Ne-
anderthals, as was suggested in Chapter 11.
Several of the peculiarities that we first saw in Saccopastore
and later in the Wiirm I Neanderthals are present in the ancient
North Africans as well as in Sinanthropus, and North Africa is
nearer to the Neanderthal home than China is.
But a principal question remains: did these North Africans sim-
ply die out, or did they evolve further into one of the five living
human subspecies? Certain archaeological, anatomical, and geo-
graphical facts support the concept that they were the ancestors
of the Bushmen. One of these is a persistent folk memory in the
oral literature of the Riffians of northern Morocco, descendants of
the Mouillians.
The Riffians have a vivid image of their predecessors, food
gatherers who would have survived longer in the inaccessible Rif-
fian mountains than on the plains below. They were, according
to legend, a people called amziw (male) and thamza (female),
and dwelt in huts built on the sides of mountains. The women
were exceedingly ugly, and their breasts dragged on the ground,
squirting milk as they walked. Their lips were long and slobber-
ing; their hair long, tangled, and curly. The men had similar lips
and hair. These people were cannibals and took delight in crunch-
ing and gnawing human bones. They had the ability to transform
themselves: a thamza could turn into a bewitchingly beautiful
602
Africa
Berber damsel, and an amziw into a Negro. Obviously, then, they
were, in their natural forms, neither Caucasoid nor Negro.9
Aside from these powers of transformation, there was nothing
supernatural about these cannibals, admittedly long extinct. They
were not concerned with jinns, shaitans, angels, gnomes, ghosts,
or other categories of spooks, haunts, and genii loci familiar
throughout most of the Islamic world. As nothing is said about
their size, it was probably normal, like that of the ancestors of the
Bushmen whose bones have been exhumed in other parts of Af-
rica.
If the Ternefine-Tangier folk were not the ancestors of the
Bushmen, they were a sixth subspecies that uniquely died with-
out modern descendants, and the Bushmen would have had no
discernible ancestors.
The Manclihle from Haua Fteah, Cyrenaica 1
So far, we have described the pre-Caucasoid fossil remains
from only the western part of North Africa, Morocco and Algeria.
Egypt has yielded no known early human remains, nor, until
1952, had Libya. In that year Charles McBurney excavated a
huge limestone solution cavity called Haua Fteah (the open cis-
tern) in Cyrenaica. He had reached the bottom of a Lower
Levalloisio-Mousterian level and was unearthing a jumble of ani-
mal bones, four feet lower down, when he found a fragment of
human mandible.
Although the industry resembled that of Tabun in Palestine, the
Carbon- 14 date of the bottom of the level in Haua Fteah was only
32,000 b.c. (No. W-85, 34,000 ± 2,800 years). McBurney inter-
polated the date of the manible at about 38,000 b.c. Both dates
fall within the time span of the Gottweig Interstadial of Europe,
much later than the supposed date of Tabun.
The fragment consists of a left side, including nearly all of the
ascending ramus, from the location of the second premolar to the
gonial angle and up to the condyle. Only the second and third
9 C. S. Coon: “Tribes of the Rif,” HAS, Vol. 9 (1931), p. 155-
1 C. B. M. McBurney, J. C. Trevor, and L. H. Wells: “The Hauah Fteah Fos-
sil Jaw,” JRAI, Vol. 83 (1953), PP- 71-85-
Earliest Caucasoid Invaders of North Africa: the Mouillians 603
molars are present, and the third molar is freshly erupted, indicat-
ing an age of eighteen to twenty-five years. Trevor and Wells con-
sider the specimen female.
This mandible is much smaller than any of the northwest Af-
rican ones in all dimensions and falls within the size range of the
Mount Carmel series, being closest to Tabun 1; but the wide angle
of its ascending ramus ( 1130 ) is closer to that of Tabun 2 ( 1180
compared to 104° for Tabun 1). Morphologically, it is difficult to
compare this mandible with others because it is badly battered.
However, the leading edge of the ascending ramus is flush with
the rear border of the third molar, and this is a non-Neanderthal
feature.
The second molar is much smaller than any of the Ternefine-
Tangier line, and the third is smaller than any but Ternefine 3,
which is abnormally short anteroposteriorly. Both Haua Fteah
teeth fit within the Mount Carmel range. The second molar is
larger than the third and neither is taurodont; the cusp pattern of
the third molar is +5.
Eastern Barbary, then, was during the Gottweig Interstadial a
refuge for a Levalloisio-Mousterian industry of an earlier Pales-
tinian type — unless the whole Mount Carmel dating is wrong. In
the latter case, the Haua Fteah industry was a contemporaneous
extension of the Tabun industry into Africa. In either case, this
evidence suggests that by the time of the Gottweig Interstadial
a presumably sapiens Caucasoid people, like the Mount Carmel
population, may have penetrated northeast Africa. These peo-
ple must have been in contact with the northwest Africans of that
period, and may perhaps have occupied the Nile Valley. If the
northwest Africans had not already become sapiens by local evo-
lution, here was their opportunity to rise to the sapiens grade
through gene flow, and to acquire a measure of Caucasoid char-
acteristics some 25,000 years before the arrival of the Mouillians.
The Earliest Caucasoid Invaders of Noii:h Africa:
the Mouillians
Before the Pleistocene was over, northwest Africa was in-
vaded by Caucasoids, contemporaries of the late Magdalenian
604
Africa
peoples of Europe. They brought with them a blade and microlith
culture called Mouillian after its type site, La Mouillah, 35 miles
west of Tlemcen in western Algeria.2
We know that Mouillians came in with a Palearctic fauna, be-
cause bones of the brown bear ( Ursus arctos ) , the Barbary sheep,
or aoudad ( Ammotragus lervius), and the Barbary ape ( Macaca
inuus ), were found in the Mouillian site of Afalou-bou-Rhummel
in Algeria. We also know roughly the date of their arrival — not
long before 10,000 b.c. — because of a Carbon- 14 dating in the
next to earliest Mouillian level in the Moroccan cave of Taforalt
(No. L-399E, 11,900 ± 240 B.P.). Because McBurney could find
no exact counterpart of the Mouillian flint industry or of their
physical type in the Near East, and because of their westerly geo-
graphical distribution, he believes that they came from Spain.3
Briggs, on other grounds, derives them from the Near East,4 and I
am inclined to agree with Briggs, for three reasons: we do not yet
know everything about the Upper Paleolithic industries of the
Near East; the earliest Mouillian skull we have is Near Eastern
Caucasoid in type, and the others could have been affected by
local mixture; and bears, aoudads, and Barbary apes could hardly
have swum across the Strait of Gibraltar, but they could easily
have walked from Palestine during' a climatically suitable period,
such as the last advance of the Wiirm.
The Mouillian culture lasted well into the post-Pleistocene pe-
riod, and its most characteristic physical type — stocky, broad-
faced, and snub-nosed — may still be seen among individual
Berbers living in relatively inaccessible regions along the Medi-
terranean coast, particularly in Kabylia and the Moroccan Rif. As
late as the time of the Spanish conquest of the Canary Islands,
during the fifteenth century a.d., some of the native Canarians,
called Guanches, especially those living on Tenerife and Gran
2 This culture was originally named Ibero-Maurusian because of its resem-
blance to a Mesolithic industry in Spain. Its name was later changed to Oranian
because of its concentration in the Department of Oran, and finally to Mouillian,
after the site in which it was first found.
3 McBurney: The Stone Age of Northern Africa (London: Penguin Books;
i960), p. 225.
4 Briggs: op. cit., p. 58.
Earliest Caucasoid Invaders of North Africa: the Mouillians 605
Canaria, were Mouillians physically, as are some of their mixed
descendants today.
The roster of Mouillian skeletons and parts of skeletons listed
on Table 32 indicates a total of more than 252 individuals. Many
of these cannot be used here, however, because they were ex-
TABLE 32
MOUILLIAN SKELETAL MATERIAL
Site Material
ALGERIA
Afalou-Bou-Rhummel, south shore
of Gulf of Bougie, Constantine
Ali Bacha, near Bougie, Constantine
Gambetta, 10 mi. SSE of Souk
Ahras, Constantine
Kef-oum-Touiza, 45 SE of B6ne,
Constantine
La Mouillah, 35 mi. W of Tlemcen,
Oran
MOROCCO
Dar es-Soltan, 4 mi. SW of Rabat
Taforalt, 33 mi. NW of Oujda, in
Beni Znassen country
32 skeletons and 5 mandibles in upper level;
1 adult male skeleton and 1 infant’s skull
in lower level
Remains of 9 individuals
Remains of 2 individuals
1 skeleton
Remains of over 15 individuals
Remains of 4 individuals
Skeletons of 96 babies, 6 adolescents,
and 80 adults
humed long ago and are lost, or else details have not been pub-
lished. The most useful are twenty-eight skeletons from Afalou-
bou-Rhummel which have been thoroughly described, and the
Taforalt series, twenty-three skulls and twenty-six mandibles of
which have recently been studied by Mile Denise Ferenbach.5
5 Briggs: op. cit.
Boule, Vallois, and R. Verneau: “Les Grottes Paleolithiques de Beni Seghoual,”
A1PH, Mem. 13 (1934), Part 2.
Vallois: “Diagrammes Sagittaux et Mensurations Individuels des Hommes Fos-
siles d’Afalou-Bou-Rhummel,” TLAB, No. 5 ( 1952).
Vallois: “Les Restes Humains de la Grotte de Dar es-Soltan,” CH, No. 11
(1952), PP- 179-202.
D. Ferenbach: “Les Restes Humains Epipaleolithiques de la Grotte de
Taforalt (Maroc oriental),” CRAS, Vol. 248 (1959), pp. 3465-7.
Ferenbach: “Les Hommes du Mesolithique d’Afrique du Nord et le Probleme
des Isolats,” BSPC, Vol. 8 (i960), pp. 1-16.
6o6
Africa
The Capstans
A second Mesolithic blade and microlith culture has been
identified in northwest Africa. This is the so-called Capsian,
named after its type site of Gafsa, about 50 miles south of Kas-
serine, of World War II fame. Its affinities are broadly Palestinian
and there is little question but that it came from the Near East
early in post-Pleistocene time.
Capsian sites fringe the Mouillian area on the east and south,
and in some sites Capsian levels overlie Mouillian deposits. But in
the northwest, particularly along the coast, the Mouillian lasted
into the Neolithic, with which both these cultures gradually
merged.
The oldest Capsian Carbon-14 date is 6,450 b.c. (8,400 ± 450
B.P., L-134), from el-Mekta, a site 10 miles northwest of Gafsa in
Tunisia. As this is an Upper Capsian site, the Lower Capsian indus-
try of the entire region must have been an earlier date, but not as
early as the Early or even Middle Mouillian.
Although the Mouillians and Capsians were both Caucasoids,
the broad-faced, heavily-muscled Mouillian type is less common
among the Capsians, who tend to be more like the original Near
TABLE 33
CAPSIAN SKELETAL MATERIAL
Site Material
TUNISIA
Ain Meterehem, 40 mi. SE of Tebessa
1 skeleton
ALGERIA
Aioun Beriche, 8 mi. N of Ain Beidha,
Constantine
More than 8 skeletons
Mechta al-Arbi, 35 mi. SW by W of
Constantine
32 skeletons, 8 skulls of which have been
described
Grotte du Cuartel
Grotte du Polygone
Grotte des Trogdolytes J
in the
• city of
Oran
Many skeletons; 3 skulls survive
MOROCCO
Sidi Ahmed el-Habib, 12 mi. W of
Berkane near the Algerian frontier
1 skeleton
The Racial Anatomy of the Mesolithic North Africans 607
Eastern prototype. As we shall see later, the Capsian culture was
carried across the Sahara into East Africa, as far south as Olduvai
and beyond, and the Capsian skeletons of East Africa bear a
family likeness to those north of the Sahara.
On Table 33 are listed more than fifty-eight Capsian skeletons,
mostly from Tunisia but represented by one site each from Algeria
and eastern Morocco. Briggs, our chief source for this material,
was able to locate and measure only eleven of the skulls, four of
which are male and seven female. He published them not in a
separate series, but in a composite North African Mesolithic series
that also included thirty-three Mouillian skulls from Afalou-bou-
Rhummel and elsewhere. In his analysis of this series by morpho-
logical types, Briggs distinguishes between the skulls of the two
cultures, which differ as previously indicated.
The Racial Anatomy of the Mesolithic North Africans
In addition to Briggs’s series, we have Vallois’s of twenty-
eight Afalou-bou-Rhummel skulls, and Ferenbach’s of twenty-
three skulls and twenty-six mandibles from Taforalt. The Mouil-
lians are far better known than the Capsians, therefore, and the
following description applies principally to the former.
The mean cranial capacity of the males is 1,614 cc- f°r a pooled
series of thirty -nine male skulls (Briggs and Ferenbach) and
1,519 cc. for seventeen female skulls. These skulls are very large,
and show considerable sexual dimorphism in their dimensions. In
metrical details the two series (Briggs’s includes the skulls pub-
lished in Vallois’s series) generally resemble the European Upper
Paleolithic crania. They have high-vaulted brain cases of variable
shape, a few of which, from Afalou-bou-Rhummel, are brachy-
cranial. Most of them have short, broad faces with low orbits and
deep, broad mandibles with everted gonial angles.
In many of the male skulls the brow ridges are heavy, but con-
centrated over the centers of the orbits with little lateral exten-
sion. Most of the chins are projecting. Mid-facial prognathism is
usually slight or absent, whereas alveolar prognathism is medium
or pronounced in about 70 per cent of tbe specimens in Briggs’s
6o8
Africa
series. The malars are large in 60 per cent of them, and the sub-
nasal fossa slight or absent in 60 per cent also. Some of the skulls
show a forward projection of the malars below the orbits, a condi-
tion similar to that seen in Mongoloids and Bushmen; and one
male skull (Afalou 40) apparently has an index of upper facial
flatness of about 8, which is very low and within the Bushman
range.
We know very little about the teeth of these people. Their
upper median incisors had been removed in childhood, as had
also, in some cases, the upper lateral and lower median incisors.
All the adult and most of adolescent teeth were extremely worn,
and only the teeth of the Dar es-Soltan skull (C-i) have been
measured. These are not remarkably large. In the mandible of this
specimen the first molar is the largest of the three; the third is the
smallest.
In general, these skulls, disproportionately drawn from two
cultures, fluctuate metrically and morphologically between two
poles. At one extreme is a long-headed Caucasoid which resem-
bles not only Combe Capelle but also several skulls of its own
geological age in western Asia, such as my Mesolithic material
from Hotu, and the Early Bronze Age skulls from Tepe Hissar in
northern Iran. At the other is a local type characterized by a num-
ber of features not seen before in combination — a broad, short
vault; a broad, short face; low orbits; a combination of a flattish
upper face with alveolar prognathism; a prominent chin; and
flaring gonial angles. Later on we shall see a more extreme version
of the same combination in Africa south of the Sahara, where it
was apparently ancestral to the living Bushmen.
The oldest known skull of whose age we can be sure, Afalou 28,
belongs to the first type, and so does the Dar es-Soltan specimen
(C-i), which presumably is as old. Five of eleven Upper Capsian
skulls fall into a generalized modern Mediterranean category,
whereas only three of twenty-eight Mouillian skulls do. This evi-
dence suggests that the modern Mediterranean element common
to North Africa today was largely if not wholly a Capsian intro-
duction.
Information on the long bones comes entirely from Vallois.6 He
6Vallois: “Les Restes Humains . . .”
Human Evolution in Africa South of the Sahara 609
measured eleven male and- eight female skeletons from Afalou-
bou-Rhummel. The male mean stature was five feet eight inches
( 173 cm., range = 162-180 cm. ) , and the female mean was five
feet five inches ( 165 cm., range = 155-180 cm.). Unless these fig-
ures are capricious because of sampling, the sexual dimorphism of
these people in terms of stature was less than that among the Up-
per Paleolithic Europeans.
Like the Upper Paleolithic Europeans, the Afalou people had
relatively long forearms and lower legs. Their hands and feet were
large. Nothing is known of the body bones of the Capsians except
for the male skeleton from Sidi Ahmed el-Habib in eastern Mo-
rocco. It is five feet four inches tall ( 162 cm. ) and the bones are
lightly built, lacking the strong muscle markings of the Mouillians
from Afalou-bou-Rhummel and Taforalt.
In sum, the racial history of northwest Africa from about 12,000
b.c. to the beginning of the Neolithic was roughly as follows. First
came a robust Near Eastern Caucasoid, along with the Palearctic
fauna. These immigrants mixed with the local Aterian folk, pro-
ducing a population with short, broad faces, flattish upper faces,
alveolar prognathism, and square jaws. While this mixture was
taking place, many of the Aterian people were being pushed
southward beyond the Sahara. Finally a Near Eastern Mediterra-
nean of modern type, like that of the Natufians, came in from the
East with the Capsian culture, and the result is, essentially, the
present-day North African Berber population.
Human Evolution in Africa South of the Sahara
Although the evidence for human evolution in North Africa
before the arrival of the Mouillians is scanty, at least it is inter-
nally consistent with the concept of the local development of a
separate human subspecies linked at the base with Sinanthropus
and at the top with the Bushmen. In the rest of Africa evidence is
just as scarce and in addition it is confusing. The confusion stems
from three facts. South Africa is a vast refuge area which drew
more than one kind of people from the East African highlands.
Several of the key skulls found in sub-Saharan Africa are dubi-
6io
Africa
ously dated. Some of them are misleading because they were hast-
ily reconstructed at the time of discovery and have never been
dismantled and redone, as, for example, Skhul 5 was refashioned
by Charles Snow. Any skull that is important enough to serve as a
document of human evolutionary history merits this treatment.
The skeletal material available, after the elimination of several
paiticularly dubious pieces, is listed in Table 34. It comes from
twenty-two sites, eight of which may be generally labelled Early
Man, five Capoid, four Caucasoid, and only three definitely
Negro.
The “Milk” Teeth from Olduvai
A yeab before the discovery of Zinjanthropus at Olduvai Gorge,
Tanganyika, Louis Leakey found two hominid milk teeth in the
deposit just above the Zinjanthropus level.7 As they were associ-
ated with early hand axes and the appropriate fauna, they were
attributed to the base of the Middle Pleistocene, in Bed II.
One is a mint-fresh, completely unworn molar. Leakey called
it a lower second milk molar. Howell suggested that it might be
an upper second milk molar, and von Koenigswald called it an up-
per left permanent second molar. According to Leakey, the sec-
ond tooth, which is badly worn, is a left lower milk canine. Von
Koenigswald called it an upper milk canine. If both were milk
teeth they could have come from the same individual, but if the
molar was a permanent and the canine a milk tooth, they probably
did not, because the milk canines are shed before the permanent
second molars erupt.
The Olduvai molar is larger than any known tooth of Homo,
milk or permanent, in both the length and breadth of its crown.
Its dimensions fall close to those of the Australopithecines from
Swartkrans, but are smaller than those of Zinjanthropus. The
tooth is longer than it is wide, as in four of the Swartkrans teeth.
In Zinjanthropus and Homo the breadth exceeds the length. The
7 L. S. B. Leakey: “A Giant Child among the Giant Animals of Olduvai,” LIN,
Vol. 232, No. 6212 (1958), pp. 1104-5.
Howell: “European and N.W. African Middle Pleistocene Hominids.”
G. H. R. von Koenigswald: “Remarks on a Fossil Human Tooth from Olduvai,
East Africa,” PKNAW, Vol. 63, No. 1 (i960), pp. 20—5.
A Possible Negro Evolutionary Line 611
crown pattern of the Olduvai tooth, with six cusps, a slight bead-
ing on the forward edge, and a fovea at each end, can be matched
in Swartkrans.
The milk canine is not distinctive morphologically. Its crown
dimensions place it within the human range if it is an upper, and
a little outside it if it is a lower. Its size dimensions are also like
those of the South African Australopithecines. This tooth, there-
fore, has little diagnostic value.
Fig. 8o The Molar from Olduvai Bed II.
At the bottom of Bed II in Olduvai Gorge,
Lewis Leakey found a molar and a canine. The
canine is a milk tooth. The molar is probably a
left upper first permanent molar. In its size,
shape, and cusp pattern it closely resembles the
corresponding teeth of Australopithecus ro-
hustus from South Africa. ( Drawing after von
Koenigswald, i960.)
In all likelihood the two teeth did not belong to the same indi-
vidual. Nothing more can be said about the canine, but the molar
is probably Australopithecine. It resembles those of the South Af-
rican robustus group closely, and differs in size and shape from
those of Zinjanthropus, whose remains were found lower down in
the same site.
Either Australopithecines coexisted with men when the bottom
of Bed II was formed; or the teeth, particularly the molar, were
eroded out of the top of Bed I and found their way into the bot-
tom of Bed II; or else the earliest men of Olduvai had bigger teeth
than those seen in the genus Homo elsewhere in the world. In my
opinion the third alternative is virtually impossible. These teeth
probably tell us nothing about early man in East Africa.
A Possible Negro Evolutionary Line
Aside from the milk teeth just described, the remaining nine
specimens or sets of specimens listed at the top of Table 34 seem
to form a sequence, although no one else to my knowledge has in-
6l2
Africa
TABLE 34
SKELETAL MATERIAL FROM AFRICA SOUTH
OF AND INCLUDING THE SAHARA
Country
Site
Date
Material
Name or Race
Tanganyika
Olduvai
Early Middle
2 milk teeth
Australopithecine(?)
Pleistocene
Tanganyika
Olduvai
Early Middle
1 calvaria
Chellian-3 Man
Pleistocene
Kenya
Kanjera
Uncertain
4 calvaria,
H. kanamensis
long bones
Cape Prov-
Saldanha Bay
Upper Pleisto-
1 calvaria,
None given
ince
cene
1 piece
mandible
N. Rhodesia
Broken Hill
Upper Pleisto-
2 individuals:
H. Rhodesiensis
cene
1 cranium,
1 maxilla,
long bones
Tanganyika
Lake Eyasi
Upper Pleisto-
Fragments
Africanthropus
cene
1 skull
njarensis
Ethiopia
Dir6 Dawa
Upper Pleisto-
Fragment
None given
cene
mandible
Transvaal
Cave of
Upper Pleisto-
Fragment
None given
Hearths
cene
mandible
Cape Prov-
Cape Flats
Late Upper or
3 individuals:
“Australoid”
ince
post-Pleisto-
2 crania,
cene
long bones
Natal
Border Cave
Post-Pleisto-
1 adult skele-
“Australoid”
cene
ton, 1 infant
Sudan
Singa
10,000-5,000
1 calvaria
H. sapiens Capoid
B.P.
Kenya
Homa Shell
Post-Pleisto-
7 skeletons
H. sapiens Capoid
Mound
cene
Transvaal
Boskop
Post-Pleisto-
1 calvaria,
H. sapiens Capoid
cene
fragment
mandible
Orange Free
Florisbad
Late Upper or
fragment cra-
H. ( africanthropus )
State
Early post-
nium, tooth
Helmei
Pleistocene
Cape Prov-
Fish Hoek
Post-Pleisto-
1 skeleton
Capoid
ince
cene
Cape Prov-
Matjies River
Post-Pleisto-
Remains 27
Capoid
ince
cene
ca. individ-
uals
Kenya
Elmenteita
Post-Pleisto-
30 skeletons
H. sapiens Caucasoid
cene
Kenya
Gamble’s
Post-Pleisto-
5 skeletons
H. sapiens Caucasoid
Cave
cene
Tanganyika
Olduvai
Post-Pleisto-
1 skeleton
H. sapiens Caucasoid
cene
A Possible Negro Evolutionary Line
613
TABLE 34 ( continued )
Country Site Date Material Name or Race
Tanganyika
Naivasha RR
Post-Pleisto-
cene
1 skeleton
H. sapiens Caucasoid
Sudan
Khartoum
Post-Pleisto-
cene
5 individuals
H. sapiens “Negroid”
Sahara
Asselar
Post-Pleisto-
cene
1 skeleton
H. sapiens “Negroid”
Mali
Kourounko-
rokale
Post-Pleisto-
cene
11 skeletons, 2
fossil mand-
ibles
H. sapiens “Negroid”
TABLE 35
THE TEETH FROM BED II OF OLDUYAI
The Molar Compared to Lower The Molar Compared to Upper
Second Milk Molars Second Permanent Molars
Robustic- Robustic-
Length Breadth
ity
Length
Breadth
ity
Olduvai
15.0
14.0
210 mm.2
Taungs
12.8
14.0
179 mm.2
Taungs
11.0
9.0
99
Sterkfon-
tein 89
14.6
15.0
219
Makapans-
Sterkfon-
gat
12.5
10.5
131
tein 20
14.0
13.2
185
Swartkrans
13.2
12.1
159
Olduvai
15.0
14.0
210
Kromdraai
12.0
9.7
116
Zinjan-
thropus
17.0
18.1/20.3
308/345
Sinanthro-
Pithecan-
pus 127
12.2
10.1
123
thropus 4
Sinanthro-
12.0
14.0
168
pus 33
12.1
13.4
162
The Canine Compared to Upper and
Lower Milk Canines
Length Breadth Robusticity
Olduvai
6.7
5.7
38 mm.2
Upper
Taungs
6.8
5.8
39
Lower
Taungs
6.5
5.3
34
Sterkfontein (largest)
6.4
5.6
36
Swartkrans (largest)
6.4
4.7
30
Kromdraai
5.2
4.9
26
terpreted them in this fashion. They begin with what is clearly a
Homo erectus brain case and end with skeletons that are primi-
tively sapiens.
In my opinion this is a Negro line, located in East and South
Africa, a line which is separated in space, and probably also in
614 Africa
time, from the three specimens at the bottom of the table, which
are indubitably Negro. In studying each specimen in turn, we
must remember that East Africa was closer to the sources of more
lapid evolutionary change in the north than South Africa was,
and that the farther south one goes, the slower the pace of human
evolution and the greater the time lag in the procession of Paleo-
lithic industries. What happened in the Sudan and West Africa,
the homelands of the modern Negroes, is still a complete mystery.
The Chellian-3 Skull from Olduvai
During his i960 excavations at Olduvai Louis Leakey found a
human skull in Bed II, 43 feet above the dividing line between
Bed I and Bed II, and in association with hand axes and cleavers
of the African Chellian industry. Because the tools at that level
belong to the third of several recognized stages, the specimen is
called, temporarily, the Chellian-3 skull of Olduvai. Like the re-
mains below it, this skull was associated with animal bones, in
this case the bones of full-sized game, which had been broken for
marrow. Chellian-3 man was apparently a full-scale hunter.
The skull consists of a faceless calvaria, broken open at the top
as well as in the base. It has very large brow ridges, a sloping fore-
head, a nuchal crest, and small mastoids: the hallmarks of the
Homo erectus grade. Its length of 209 mm. is excessive, and its
breadth of about 133 mm. is narrow, for an otherwise large erec-
tus skull. In fact, this breadth dimension would fit the intercondy-
lar breadth of the Heidelberg jaw. Its probable auricular vault
height of about 109 mm. is low. Its walls are apparently thick.
Probably its cranial capacity was between 1,100 and 1,200 cc.,
nearly equivalent to the capacities of the largest Solo and Sinan-
thropus skulls.8
However, it differed from both morphologically. The brow
ridges form a double arch when seen from in front and sweep far
to the rear on either side. Although very much larger and longer,
these brow ridges resemble those of Steinheim in details of form.
8 Only the length and breadth measurements have been published. The other
figures are my own, derived from a scale photograph and calculations.
FOSSIL MAN SITES IN AFRICA
south of and in the Sahara
i stoetor,*\ \ .
I * i Tchad . 1
Wii'mnkoroka le ' Australopitheclne-
Khartum
singa /
,/ \ \ a Direvawa
// /
Ip V ’ / Oloraesaitie^ Jjp
(ARCHAEOLOGY ONLY)
, s' ^ a'i--\Amkuru
Lake Eyasi *j --dm (Nairobi)
1 Olduvat |J\ Nairn ha
— .3, fj ilmmteita
Kalambo Tails % m Gamble's Cave
(FIRST FIRE IN AFRICA) ft
* i Hearths • \ ip
^^Jtimsb*d.._ *WrderCave
Saldanha Bayk # sT
Fish HoekkL&s^'t,-' vil,ar.
Cape Flats ™atl,es ^mr
MAP 12
6i6
Africa
The index of upper facial flatness, impossible to calculate accu-
rately from present evidence, probably fell within the Caucasoid
and Negro ranges. This contemporary of Heidelberg and Sinan-
B
SALDANHA
A
CHELLIAN 3
Fig. 81 Profiles: Chellian 3, S a 1 da n ha. and Broken Hill. Of the five lines
of human descent, the two African ones are the most poorly represented by fossil
specimens. The oldest of what seems to be the Congoid line is the Chellian 3 skull
from Olduvai, shown here in a restored form. The top of the skull is missing but
enough is left of the parietals to make a fair reconstruction. Although its brow
ridges are heavy, its nuchal area is nearly modern in form. The Saldanha skullcap,
presumably over 300,000 years younger, is essentially the same; and the Broken
Hill skull, younger still, shows little advance over its predecessors. Unless the dating
of the last two skulls is wrong, human evolution proceeded at a snail’s pace during
the Middle and Upper Pleistocene in Africa south of the Sahara. ( Drawing A after
Leakey; B after a photograph by the author; C after Pycraft. )
thropus (Leakey, Evernden, and Curtis set the date at 360,000
years by Argon-40 analysis) 8 could have been close to an even
earlier common ancestor of both Caucasoids and Congoids.
9 Leakey, J. F. Evemden, and G. H. Curtis: “Age of Bed I, Olduvai Gorge,
Tanganyika,” Nature, Vol. 191, No. 4787 (1961), pp. 478-9.
The Kanjera Specimens
617
It is tempting also to relate Chellian-3 man to its local prede-
cessor, Zinjanthropus. But in at least two respects Chellian-3 is
more primitively hominid, or even pongid, than Zinjanthropus, or
indeed any other known Australopithecine. Its nuchal crest, like
that of Pithecanthropus 4, sits higher on its occipital bone, and its
foramen magnum lies farther to the rear in the base of the skull.
To derive Chellian-3 man from Zinjanthropus would be biologi-
cally impossible.
We must not let ourselves be misled into interpreting the con-
tinuity of stone implements in Beds I and II of Olduvai Gorge to
indicate a continuity of tool makers — from Zinjanthropus to Chel-
lian-3 man. The crude tools that both beds have in common were
of a type made all the way from Morocco to South Africa and from
Palestine to Indonesia by several kinds of hominids, including,
no doubt, both Australopithecines and men. But in Bed II, for the
first time in the Gorge, hand axes appear, and they are the hall-
mark of Western man. These hand axes are few in number, com-
pared to the cruder tools, and less skillfully fashioned than those
made in Europe at the same time. What evidence there is suggests
that Chellian-3’s ancestors had come from farther north in Africa
and had not evolved from local, East African Australopithecines.
The Kanjera Specimens 1
I n 1932, the same year in which Louis Leakey found the Kanam
mandible ( see Chapter 7 ) , he also found human remains at Kan-
jera, a neighboring site on the south shore of the Gulf of Kavi-
rondo, Lake Victoria Nyanza. Kanjera is the type site of the Kan-
jeran Pluvial period, the third in the East African sequence, be-
lieved to correspond roughly to the Riss glaciation in Europe.
Three of the four specimens had weathered out and were lying
on the surface, but one of them, Kanjera 3, was still partly im-
bedded in the ground. All were covered with a crust of the gray-
ish, calcified sand in which Number 3 had lain. All four were
equally mineralized, and had the same fluorine content as the
1 Leakey: The Stone Age Races of Kenya (Oxford: Oxford University Press;
1935).
6i8
Africa
animal bones that accompanied them.2 They were also appar-
ently, but not positively, associated with late Acheulian hand
axes. We do not know the exact age of the Kanjeran fauna, nor do
we know when hand axes ceased to be made in Kenya. If, as now
seems likely, the human specimens belong to the geological setting
to which they have been attributed, they are probably of Upper
Pleistocene date. But they could be only 40,000 to 30,000 years
old. Or they could be of an earlier date, or possibly intrusive. No
one really knows, which is unfortunate because they are anatomi-
cally unique.
Number 1, consisting of seven fragments, covers most of the
sagittal profile of the brain case, except for its base, and also in-
cludes pieces of a zygomatic bone and maxilla. Number 2 is rep-
resented by three small pieces of parietal. Number 3, like Num-
ber 1, consists of seven pieces of vault, although they are less com-
plete and the bones do not all articulate. Number 4 is a piece of
frontal bone with nasion intact, and a smaller piece of vault.
The first three specimens are thick-walled, the fourth thin.
Numbers 1 and 3 are long, narrow, and low-vaulted,3 with deeply
curved frontal and occipital bones and a flat profile on top. Num-
bers 1 and 3 lack brow ridges and have a general infantile ap-
pearance whereas Number 4, the thin bone, has moderately
strong brow ridges. Inside the frontal of Number 1 an extensive
sagittal crest served as an internal brace. The left frontal bone of
Number 3 has a high temporal crest, indicating a heavy jaw mus-
culature. In Numbers 1 and 4 nasion is placed high and the nasal
bones could not have been excessively broad, at least at the root
of the nose. The zygomatic and maxillary fragments of Number 1
are small and slight, with a canine fossa. The endocranial cast
shows an enlarged area striata (the visual region of the occipital
lobes ) and an ill-filled parietal region, as in the Homo erectus
specimens from Java and China and in some modern Australian
aboriginal skulls.
The capacities of Numbers 1 and 3 have been estimated at
U350 to M°° cc-» and these figures, along with the morphology
K. F. Oakley: Physical Anthropology in the British Museum (London, 1958).
2 The reconstructed length and breadth dimensions of Number 1 are 207 mm.
and 137 mm.; the cranial index is only 66. The comparable figures for Number 3
are 208 mm., 140 mm., and 67 mm.
The Saldanha Bay Skullcap 619
of the skulls, mark them as sapiens. Without more of the faces and
with no teeth available for study, it is difficult to give them a racial
designation, but they fit the category of Negro more than any
other. A perfectly modern toe phalanx associated with Number 3,
three pieces of rib belonging apparently to Number 2, and a fe-
mur shaft found near Number 3 are all modem, although ra-
cially undiagnostic.
Whatever their date, these specimens are a primitive grade of
Homo sapiens, and are probably Negro. As the femur was full-
sized, they were presumably not Pygmies.
The Saldanha Bay Skullcap 4
The Saldanha Bay skullcap was discovered in 1953 by Ronald
Singer and Keith Jolly in a huge bed of fossil animal bones at
Elandsfontein Farm, 15 miles east of Saldanha Bay and about 90
miles north of Capetown. Along with it were found many stone
implements, including both hand axes and flake tools. A piece of
mandible was later found in the same site and attributed to the
same individual.
The date of the skull has not yet been firmly established. On the
4 M. R. Drennan: “A Preliminary Note on the Saldanha Skull,” SAJS, Vol. 50,
No. 1 (1953), PP- 7-ii-
Drennan: “The Saldanha Skull and its Associations,” Nature, Vol. 172, No.
4383 ( 1953), PP- 791-3-
Drennan: “Saldanha Man and his Associations,” AA, Vol. 56, No. 5 (1954),
PP- 879-84.
R. Singer: “The Saldanha Skull from Hopefield, South Africa,” AJPA, Vol. 12,
No. 3 (1954), PP- 345-62.
Drennan and Singer: “A Mandibular Fragment, Probably of the Saldanha
Skull,” Nature, Vol. 175, No. 4452 (1955), PP- 364-5-
A. Mabutt: “Geomorphology, Archaeology, and Anthropology from Bok Baii,
Darling District, Cape Province I, Physiography and Surface Deposits,” SAAB,
Vol. 10, No. 39 (1955), PP- 85-6.
Singer: “Man and Mammals in South Africa,” /PS/, Vol. 1 (1956), pp. 122-30.
Singer: “Investigations at the Hopefield Site,” FTP A, 1957, pp. 175-82.
Singer: “The Rhodesian, Florisbad, and Saldanha Skulls,” NC, 1958, pp. 52-62.
Singer: “The New Fossil Sites at Langebaanweg (South Africa),” CA, Vol. 2,
No. 4 (1961), pp. 385-7-
Singer and J. R. Crawford: “Archaeological Discoveries at Hopefield,” JRAI,
Vol. 88, Part I (1958), pp. 11-19.
Oakley: “Dating the Stages of Hominid Evolution,” The Leech, Vol. 28, Nos.
3, 4, 5 (1958), pp- 112-15.
620
Africa
basis of faunal associations, confirmed by fluorine tests, it could
be 100,000 years old, but on the basis of associated artifacts it
could be no more than 40,000 years old.5 For present purposes
it makes little difference which of these dates is correct, because
Saldanha man lived, in either case, between the dates of Chel-
liaii-3 man and Rhodesian man, and forms a close anatomical
link with them.
The Saldanha Bay specimen is a skullcap reconstructed from
twenty-seven pieces and completely lacking a base except for two
triangular fragments of the occipital squama, situated on either
side of the center line. Although the cranial capacity cannot be
measured exactly, Singer places it between 1,200 and 1,250 cc., a
little over the estimated figure for the Chellian-3 skullcap, which
it closely resembles morphologically. In grade it is also equivalent
to the larger skullcaps from Solo and Choukoutien, and is a classic
example of Homo erectus.
Its sagittal profile is very much like that of Chellian-3, except
that its brow ridges and occipital region are a little less protuber-
ant. It is somewhat shorter (200 mm. as compared to 209 mm.)
and probably broader (i44(?) mm. as compared to 134(F)
mm.), but the vault height above the glabella-opisthion line is ap-
parently the same in both specimens (84 mm.). Like those of
Chellian-3, its brow ridges sweep back laterally, and its index of
upper facial flatness, about 20, is somewhat high for modern Ne-
groes but normal for Caucasoids.
The piece of mandible consists of the upper forward half of the
ascending ramus, including the coracoid process and the nutrient
foramen. As reconstructed by Singer, it extends onto the body of
the right condyle and down to within 10 mm. of the lower margin
5 Singer (1961) has equated the fauna at Hopefield and that of the upper of
two levels at a very rich nearby site, Langebaansweg, which is Late Middle to
Early Upper Pleistocene. On the other hand, the accompanying Fauresmith arti-
facts, which include both very late hand axes and Middle Stone Age flakes, seem
to indicate a later date. According to Oakley ( 1958 ) , the implements represent a
transition from one industry to another rather than a sequence of two distinct
cultures. This particular local form of the Fauresmith culture has been assigned
the same general date as a forest-culture called Sangoan, which was given a
Carbon-14 date of about 40,000 years ago. The Lamont Laboratory has produced
one date of 41,000 ± 3,300 B.c. (L-399 C), which, it advises, should be used with
caution.
The Broken Hill or Rhodesian Specimens
621
of the jawbone. The ascending ramus was therefore about 57 mm.
high and 47 mm. wide. It was not high, and it rose at nearly a
right angle from the body of the mandible. The coracoid process
slopes forward, as in Heidelberg, rather than backward, as in Tern-
efine and Sinanthropus. In general, it closely resembled the cor-
responding portion of the Heidelberg jaw, but it is thinner by
1 mm. to 4 mm. at all points that can be measured.
The Saldanha Bay skullcap and mandible patently come from
the same line as Chellian-3, although the Saldanha Bay specimen
is several hundreds of thousands of years younger. This line re-
sembles the Caucasoid much more closely than it does the Austra-
loid, Mongoloid, or Capoid. The degree of its resemblance to the
Congoids will be seen once we have examined other specimens
that still have facial bones, particularly that of the Broken Hill
skull, otherwise known as Rhodesian man.
The Broken Hill or Rhodesian Specimens 6
The Broken Hill skull, also known as Homo rhodesiensis, or Rho-
desian man, was discovered in 1921 in the course of mining opera-
6 F. A. Bather, W. P. Pycraft, et al: Rhodesian Man and Associated Remains
(London: British Museum [Natural History]; 1958), including:
W. P. Pycraft: “Description of the Human Remains,” pp. 1—51.
G. Eliot Smith: “Endocranial Cast,” pp. 55-8.
M. Yearsley: “Pathology of the Left Temporal Bone,” pp. 59-63.
J. T. Carter: “Teeth of Rhodesian Man,” pp. 64-5.
R. A. Smith: “Associated Stone Implements,” pp. 66-9.
A. T. Hopwood: “Mammalia,” pp. 70-3.
D. M. A. Bate: “Aves,” p. 74.
W. E. Swinton: “Reptilia,” p. 75.
A. Hrdlicka: “The Skeletal Remains of Early Man,” SMC, Vol. 83 (1930), pp.
98-144.
F. Weidenreich: “The Skull of Sinanthropus pekinensis”; “Some Particulars of
Skull . . .”
J. D. Clark, et al: “New Studies on Rhodesian Man,” JRAI, Vol. 77, Pt. 1
(1947), PP- 7-32.
Clark: “Further Excavations at Broken Hill, N. Rhodesia,” JRAI, Vol. 89, Pt. 2
(i960), pp. 201-32.
Oakley: New Evidence Regarding Rhodesian ( Broken Hill) Man (New York:
Paper read at Viking Foundation Conference; June 20, 1950).
Oakley: “The Dating of the Broken Hill, Florisbad, and Saldanha Skulls,”
PTPA, 1955, pp. 76-9.
Oakley: “Physical Anthropology in the British Museum.”
Singer: “The Rhodesian, Florisbad, and Saldanha Skulls.”
622
Africa
tions at the Broken Hill Mine, Northern Rhodesia. The mine then
consisted of two kopjes or hills composed of lead and zinc ores.
The mining procedure was simply to break up the rock and carry
it to the smelters. In 1907 the miners cut a hole through Kopje 1
and found inside it a pocket full of animal bones. As these were
highly mineralized, they were smelted. Only in 1921 did the min-
ers notice human bones among them, and some of these were
saved.
The principal specimen is a virtually complete cranium that
came from the very bottom of the deep pit at the end of the cave.
In the same general area one tibia and one clavicle were also
found, but the clavicle was later lost. Higher up in the main shaft
and some distance away were found one broken os coxae or pelvic
bone, one sacrum, pieces of two femora, the distal end of a tibia,
and the distal half of a humerus, as well as a piece of a second
maxilla with the right second and third molars in place. There is
no assurance that any of these bones go together, even in the case
of the tibia found with the skull, but they are all of roughly the
same age.7
The associated artifacts are flake tools of an industry known as
Proto-Stillbay to Stillbay, one of a series of African flake cultures
of the so-called Middle Stone Age. Similar tools were found by
Desmond Clark and his associates in a trench dug in the front of
the kopje. They come from the end of the Lower Gamblian Pluvial
period, about 23,000 b.c.8
The associated fauna contains only species still in existence,
with two exceptions; a rhinoceros ( Diceros whitei Chubb), and a
serval cat ( Leptailurus hintoni ), both of which could have be-
come extinct only recently.
The late Ales Hrdlicka, who was given neither to overstatement
7 At first it was believed that the skull and the other bones were of different
ages because the skull was found in lead ore and the bones in zinc ore. Subse-
quent analysis showed that both skull and bones were impregnated with zinc,
and therefore the mineral in which each lay simply reflected its eventual position
in the cave ( Oakley, 1950 ) . Still further analysis indicated that the nitrogen con-
tent of the skull and that of the bones are the same, and so is the age of both
(Oakley, 1955).
8 J. D. Clark (1947, i960) determined this date by comparing the finds in the
trench with the previously established sequence at another Rhodesian site,
Mumbwa Cave.
The Broken Hill or Rhodesian Specimens 623
nor to displays of unbridled emotion, called the Broken Hill skull
“a comet of man’s prehistory.” 9 In 1930, when he made this pro-
nouncement, Solo, Saldanha, Zinjanthropus, and Chellian-3 had
not yet been discovered and Sinanthropus was being chiseled
from its breccia. Thirty years later, the Broken Hill skull is still un-
usual in appearance, probably because it is the only complete
skull of its evolutionary grade that we have. Also, it shows an in-
congruous combination of archaic and modern, brutal and deli-
cate features, which label it a tired-looking peripheral survivor of
an ancient and vigorous race of very primitive men.
In general morphology, Broken Hill closely resembles both
Chellian-3 and Saldanha. It is as long as Chellian-3 and as broad
as Saldanha. Its auricular height was less than Chellian-3’s
(105 as compared with ca. 109 mm.), but its projected height
above the glabello-opisthion line was greater (85 as compared
with 79 ca. mm.) The reason for these contradictions in the two
height measurements is that in the Broken Hill skull, as in Sal-
danha, the nuchal crest is set lower on the occipital bone than in
Chellian-3.
The cranial capacity of the Rhodesian skull is 1,280 cc., a little
more than that of any other Homo erectus specimen we have yet
seen, if the reconstructed figures for fragmentary skulls are cor-
rect. Its internal dimensions 1 match Solo’s; the Rhodesian skull,
however, is internally higher, but by only about 4 mm.
As the skull had been protected in the cave, it was not weath-
ered or battered internally or externally. Because nearly all the
right side of the base had been broken off at the time of discovery,
Mr. Barlow of the British Museum, a peerless technician, was
able to make a perfect endocranial cast, which was studied by the
distinguished brain specialist Sir Eliot Smith.
Sir Eliot found that the track of the middle meningeal artery
was simple, as in Sinanthropus, Solo, and Ternefine, but that its
anterior branch was large. There is a large Sylvian crest, and the
prefrontal, upper parietal, and lower temporal areas of the cortex
were poorly developed in comparison with those of living men
and of the Neanderthals. The imprint of the basal surface of the
9Hrdlicka: “The Skeletal Remains . . . ,” p. 130.
1 Length is 173 mm.; breadth is 136.5 mm.; and height is 114 mm.
624 Africa
cerebellum shows that its lobes were more rounded and bent far-
ther downward than in Solo.
The imprint of the pituitary fossa indicates a sella turcica of
about the same size as that of Solo 11 (20 by 20 mm.). As the
basal portion of the sphenoid bone was in perfect condition and
the sella turcica easily visible, neither the shape nor the size of
the seat of the pituitary is open to question.
The bones of the cranial vault are thinner than Saldanha’s, the
thickness of the right parietal ranging only from 6 to 10 mm. Next
to Chellian-3, the Rhodesian specimen has the broadest, most
Fig. 82 Profiles: a Possible Negro Line— Broken Hill, Cape Flats, and
Border Cave. The modern Negroes seem to have evolved in West Africa and the
Sudan, where we have no Negro skulls of Pleistocene age. In East and South
Africa a number of skulls have been found which are apparently of early post-
Pleistocene date. Of these the most authentic seem to be the Cape Flats and
Border Cave skulls. These two are apparently earlier than the first Bushman re-
mains in those regions, and they may provide a continuity between the Broken Hill
skull, which was morphologically Congoid, and a primitive Negro population which
preceded the ancestors of the Bushmen in South Africa. (Drawing A after Pycraft
1928; B after A. Keith, 1931; C after Cooke, Malan, and Wells, 1945 and 1950. )
massive brow ridges of any human skull yet found; they match
those of Zinjanthropus.2 In form the Rhodesian brow ridges
closely resemble those of Chellian-3 and Saldanha, sweeping far
backward to either side.
Refore we study the most remarkable part of the Rhodesian
skull, its face, let us consider the teeth. The right lateral incisor
had been lost during the individual’s lifetime; both right premo-
lars, the left second premolar, the left first molar, and the third
2 The width and thickness of the brow ridges of Chellian-3 are, tentatively,
147 by 25 mm.; of Saldanha, 122 by 21 mm.; of Rhodesian man, 140 by 21 mm.;
and of Zinjanthropus ca. 140 by ca. 20 mm.
The Broken Hill or Rhodesian Specimens 625
right molar had been reduced by a combination of caries and at-
trition to mere shells, and the right canine had been worn or
broken to a stub. (For the dimensions of the other nine teeth, see
Table 39.)
The measurable teeth are as large as those of Sinanthropus, and
one of them, the right upper second molar, is larger than the cor-
responding tooth of Pithecanthropus 4. The Rhodesian teeth are
also larger than the three upper teeth from Tangier. The Rhode-
sian front teeth, from the upper lateral incisors to the first upper
premolars, are as large as those of Zinjanthropus or larger; the re-
maining teeth, from the second upper premolar to the third molar,
are smaller. The length of the outer dental arc, from third molar to
third molar, is 175 mm. in the Rhodesian skull and 195 mm. in
Zinjanthropus. The dental equipment of Rhodesian man was as
massive as that of any other Homo erectus.
As in Sinanthropus and most modern individuals, the first mo-
lars were the largest and the third molars the smallest. The Rho-
desian molars were taurodont. There is no evidence of a cingu-
lum, Carabelli’s cusp, or other unusual excrescences, and as far as
we can tell the incisors were not markedly shoveled, if they were
shoveled at all. A Flower’s index of 44.2 makes Rhodesian man
megadont. In general, these teeth resemble those of modern Ne-
groes more than those of any other race.
As in Zinjanthropus and the Neanderthals, the palate of the
Rhodesian skull is perched far forward of the middle portion of
the skull base. Although this position indicates a considerable
total facial prognathism, the face itself is so long that the principal
inclination of the upper jaw is downward. The upper face height,
93 mm., is the greatest recorded for any fossil skull, except for the
Neanderthal La Ferrassie male, whose face may have been 3 mm.
longer. Weidenreich’s reconstruction of Pithecanthropus 4 has
an upper face height of only 89 mm.
On the other hand, Rhodesian man’s bizygomatic diameter of
148 mm. is not remarkable. The same figure is found for the Cro-
Magnon skull. Rhodesian man did not need flaring zygomatic
arches because the forward position of his face left enough room
behind his eye sockets to accommodate his temporal muscle bun-
dles, despite heavy chewing. This does not mean that his orbits
1
626
Africa
were shallow like those of Sinanthropus. They were high, wide,
and deep — as far as I know the largest orbits yet found in Homo.
Like the Neanderthals he had a puffy facial surface, without
canine fossae. In the frontal index of facial flatness and in the
simotic index (reflecting the arching of the nasal bones at their
root), the Rhodesian skull falls within the Caucasoid range, and
in the third or rhinial index of facial flatness it resembles the an-
cient Egyptians and approaches the means of published series of
living Negroes. In the fourth or premaxillar index of facial flatness
it leaves all modern populations far behind it. In other words, this
face is Caucasoid in its upper portion, Congoid in the middle,
and virtually pongid below. On the whole this face is mostly
Negro.
The tibia, which was found in the bottom of the cave with the
skull, resembles that of a modern Negro in all essential details,
and indicates a stature of about five feet seven inches ( 169.5 cm. ) •
The palate of a second individual, recovered from the front of
the cave, consists of the lower half of the right maxillary bone,
with the posterior half of the right second molar and the entire
right third molar still embedded in it. The outer surface of this
piece of bone has been scraped off except for that portion imme-
diately over the third molar. The maxillary height, from the base
of the nasal cavity to prosthion, was 24 mm., compared to 35 mm.
for the skull. This difference indicates either that the Rhodesian
male population was variable in upper jaw height, that this palatal
fragment belonged to a female, or that this palate may not have
been that of a member of the Rhodesian population. Its third mo-
lar is even larger than that of the corresponding tooth in the Rho-
desian skull, which it resembles in form.3
In the debris from the front of the cave were also found two
fragments of pelvic bone, a right and a left, which belonged to
two individuals. The right one is modern in type whereas the left
one has a number of peculiarities, including a high ilium and a
shallow acetabulum, which led Pycraft 4 to assign not only this
bone but also Rhodesian man as a whole to a new genus, CypJian-
thropus rhodesiensis Woodward (Stooping Man). This designa-
3 Length is 10 mm.; breadth, 12 mm.
4 Pycraft: Description of the Human Remains.
The Cranial Fragments from Lake Eyasi, Tanganyika 627
tion has been ignored by subsequent authors. Also available are a
small sacrum, the lower portion of a large humerus, and parts of
two femora from different individuals. None of these bones devi-
ate conspicuously from those of modern Negroes.
The Cranial Fragments from Lake Eyasi, Tanganyika 5
Another specimen from East Africa probably belongs to the
same category as Rhodesian man, if it can be given any taxo-
nomic position. It is the so-called Africanthropus njarensis dis-
covered by Ludwig Kohl-Larsen on the shore of Lake Tangan-
yika, in 1935 and 1938.6
Eyasi man, as this specimen may be called, consisted of over
two hundred scraps of skull, probably belonging to as many as
three individuals, which Weinert painstakingly fitted together, fi-
nally producing a complete brain case. It was lost when Kiel was
bombed during World War II, and we have only Weidenreich’s
estimate 7 that it was probably a member of the Rhodesian popu-
lation and a female. Animal bones and stone tools were found on
the surface with the skull fragments. Rut as chemical tests of the
human bones cannot be made, the specimen cannot be dated.
The Mandibular Fragment from Dire Daiva, Ethiopia 8
I n xg23 the dean of French prehistorians, the late Abbe Henri
Rreuil, found a piece of human mandible in the so-called Porcu-
pine Cave in the hill of Balia, two kilometers from the Ethiopian
city of Dire Dawa. Although the floor of the cave contained a
number of industries, the mandible, which was fossilized, was
5 H. Weinert, W. Bauermeister, and A. Remane: “Africanthropus njarensis,
Beschreibung und phyletische Einordnung des ersten Affenmenschen aus Osta-
frika,” ZfMuA, Vol. 38 (1940), pp. 252-308.
6 The generic name Africanthropus was subsequently dropped because it had
already been assigned, equally without justification, to the Florisbad cranial
fragment.
7 Weidenreich: “The Skull of Sinanthropus pekinensis,” p. 221.
8 H. Breuil, P. Teilhard de Chardin, and P. Wemert: “Le Paleolithique du
Harrar,” L’Anth, Vol. 55, No. 3-4 ( 1951 ), pp. 219-30.
Vallois: “La Mandibule Humaine Fossile de la Grotte du Port-Epic pres Dire-
Dauoa ( Abyssinie),” L’Anth, Vol. 55, No. 3-4 ( 1951), pp. 231-8.
6z8
Africa
probably of the same age as the Stillbay implements; these are
similar to those found at Broken Hill.
The mandible consists of a piece of the right branch extending
from the first premolar past the third molar, and lacking both sym-
physis and ascending ramus. Enough is left, however, for us to see
that it was a large jaw, in the size range of Heidelberg, the North
African mandibles of the Ternefine-Tangier line, the larger Sinan-
thropus mandibles, and those of some of the Neanderthals.9
The teeth are in poor shape, with most of the enamel and den-
tine gone from the crowns. Their measurements, published by
Vallois,1 are individually within the modern range and smaller
than those of the Broken Hill skull.2 Yet Vallois finds a total length
of the premolar-molar row, from the front of the first premolar to
the back of the third molar, of 52 mm., the same as for Heidel-
berg, and more than the figure for Broken Hill.
Although this mandible is classified in the literature as that of a
Neanderthal, I see no justification for this label, nor can I find any
clue to its actual relationship. It is included here in the Chellian-3
to Rhodesian group solely because of its geographical and cul-
tural position.
The Mandible from the Cave of Hearths 3
I n the Cave of Hearths at Makapansgat, Transvaal, the scene of
Australopithecine discoveries, Raymond Dart found, in 1947, a
human mandible cemented in red breccia and associated with the
9 At the level of the mental foramen its height is 34.0 mm., its thickness 16.3
mm., and its robusticity index 47.9.
1 Vallois: “La Mandibule . . . de Dire Dawa.”
2 The Dire Dawa tooth measurements are:
First
Second
First
Second
Third
Premolar
Premolar
Molar
Molar
Molar
Mesiodistal
5
5.5
9.9
10.5
10 (?) mm.
Labiolingual
7
6.5
9
10
9 mm.
Robusticity
35
36
89
105
90 (?) mm.2
3 R. A. Dart:
“The First
Human Mandible from
the Cave
of Hearths,
Makapansgat,” SAAB, Vol. 3, No. 12 ( 1948), pp. 96-8.
P. V. Tobias: “The Kanam Jaw,” Nature, Vol. 185, No. 47 14 (i960), pp.
946-7-
Tobias: “Early Members of the Genus Homo in Africa,” in G. Kurth, ed.:
Evolution und Hominisation, Festschrift zum 60. Geburtstag von D. G. Heberer
(Stuttgart: G. Fischer Verlag; 1962).
The Mandible from the Cave of Hearths 629
final type of hand axes known in South Africa. It probably dates
to about 40,000 years ago, plus or minus 10,000 years,4 and it was
certainly no more recent than the Broken Hill skull.
It consists of a piece of the symphysis and right body of a man-
dible, running from the socket of the lower left median incisor to
the place where the right lower third molar would have been had
it erupted. Only the first premolar, first molar, and second molar
are preserved, the other teeth being represented by broken roots,
or having been lost. As the first molar was worn and the second
molar mint-fresh and crisp-cusped, Dart gives the specimen’s age
as twelve.
It is a stout little mandible, probably chinless, with an angle of
inclination of about 62°, which places it in the same class as Ter-
nefine 1, Sinanthropus G, and Heidelberg.5 Morphologically it
shows no distinctive features. It has only one foramen on its re-
maining side, and no torus mandibularis.
The three teeth are too small to match the upper teeth of Broken
Hill man, and smaller than those of the Ternefine-Tangier folk of
North Africa; their closest counterparts are in the Heidelberg
jaw.6 The first lower premolar is pointed and looks like a canine,
with a very low buccal cusp. The molars are plain, without wrin-
kles or cingulums, and the second molar has a standard Y-5 cusp
pattern.
This mandible and its teeth probably belong to the Chellian-3
to Rhodesian group. However, if I were to see it without knowing
where it came from, I would probably guess that it was of Euro-
pean origin, and set the time at the Third Interglacial.
4 I arrive at this general date by comparison with the known dates of other
sites in East and South Africa. In the Cave of Hearths itself a Middle Stone Age
level was given a Carbon-14 date of 15,111 ± 730 B.P. (C-925). Libby gives
Middle Stone Age III a date of 9,650 ± 700 ( C-924 ) .
5 At the level of the first molar the height is 25.5 mm., the thickness 16.0 mm.;
the index of robusticity 47.9. The indices of robusticity are 54.5 for the symphy-
seal region and 46.4 for the region of the mental foramen. The bone constituting
the chin region had been scraped off.
First
First
Second
Premolar
Molar
Molar
Length
8.0
12.0
11.5
mm.
Breadth
8.0
11.8
10.7
mm.
Robusticity
64
142
123
mm.2
630
Africa
The Cape Flats Skull
The line that we have tried to trace, from Kenya to Cape-
town and from the Early Middle Pleistocene to almost the end of
the Upper Pleistocene, must have survived into the Recent period.
Very likely it evolved eventually from a subspecies of Homo erec-
tus into a subspecies of Homo sapiens, either by a purely local evo-
lutionary process or by gene flow from outside. It may even have
been wholly absorbed into another line. I think that we can rule
out independent mutation as the cause of change, because the
territory inhabited by this ancient line is fully exposed to contacts
from the north.
Because South Africa is the most remote and isolated part of the
continent — next to the rain forest, from which no pertinent infor-
mation is available — it is there that we may expect to find survi-
vors of the Chellian-3 to Rhodesian line in post-Pleistocene time.
During the last half century a number of local archaeologists,
both amateur and professional, have found hundreds of buried
skeletons, mostly in rock shelters, which date from shortly after
the end of the Pleistocene to the time when Dutch gin bottles ap-
pear amid the cultural debris, and after. Most of these finds are
difficult to date, and some are actually post-Dutch, but among
them it may be possible to locate what we are looking for.
This search is not helped by the fact that, except for Ronald
Singer, apparently everyone who has worked with this material
assumes that all skeletons found in South Africa, whatever their
antiquity, must in some way be ancestral to the living Bushmen,
Hottentots, and Korana (a kind of Hottentot), all of whom, in
their opinion, were autochthonous.7 However, there is an alterna-
tive theory of the origin of the Capoids: that they came from the
north in postglacial times. With this in mind, we can examine
each of the postglacial South African skulls critically to see if some
of them are not only early in date but also racially non-Capoid. A
number of non-Capoid skulls are indeed available. They have
7 Singer: “The Boskop ‘Race’ Problem,” MAN, Vol. 58, Art. No. 232 (1958),
PP- 1-5-
The Cape Flats Skull 631
been set aside by the South African anthropologists and classified
as “Australoid” or “gerontomorphic.” Two of them seem to be of
moderate antiquity; these are the skulls from Cape Flats and the
Border Cave.
In 1929 M. L. Drennan discovered an open-air site at Cape
Flats, near Capetown, which contained, at a depth of three feet,
the remains of three individuals, one skull of which has been
partly described.8 It was found without fauna in a deposit con-
taining Stillbay flake tools, but there were also Wilton implements
in the neighborhood. Stillbay tools are the kind associated with
Broken Hill, and Wilton is a local Capsian derivative made by
Bushmen. In South Africa the Middle Stone Age continued as late
as 5,000 years ago,9 and the Cape Flats site could have been, but
was not necessarily, of that late.
The Cape Flats skull is long and narrow, with a cranial capacity
of 1,230 cc., the same as Saldanha’s and slightly less than Broken
Hill’s, and its vault is but two millimeters higher than the latter’s.
According to Drennan, the endocranial cast reveals the same
primitive features seen in the Broken Hill skull. So far it indicates
little or no advance in the shape of the brain of the line to which
it seems to belong. But the external morphology of the brain case
is more modern.
The brow ridges, although heavy and sweeping backward, are
smaller than those of the earlier skulls. The brain case is angular,
but in contour it is modern. Indeed, it generally resembles some
Australian skulls, but it also looks somewhat like Steinheim. The
face is long but not in the same class as the Broken Hill face, and
in the degree of facial flatness, which cannot be measured without
access to the original specimen, it follows the Rhodesian pattern
of combining Caucasoid and Negro. The lower jaw has a chin.
Unfortunately no incisor teeth were preserved in either jaw.
There are five upper teeth, from canine to first molar, and six low-
ers, from canine to third molar. The upper teeth are smaller than
8 Drennan: “An Australoid Skull from Cape Flats,” JRAI, Vol. 59 (1929),
p. 417.
A. Keith: New Discoveries Relating to the Antiquity of Man (London: Wil-
liams & Norgate; 1931), pp. 140-2.
9 A Middle Stone Age date from the Holley II site is 4,490 ± 150 B.P. (BM-34).
632 Africa
those in the Broken Hill skull, and the lowers match those from the
mandible of the Cave of Hearths. All fall within the modern Ne-
gro range.
As Drennan pointed out, this skull does not resemble that of a
modem South African Bantu, but there is no reason for it to do
so. The Bantu are as recent as the Boers in South Africa, and this
skull may well be old enough to have antedated the arrival of the
ancestors of the Bushmen. If that is the case, then this skull may
represent a descendant of the Chellian-3 to Broken Hill line which
had crossed the sapiens threshold but had not evolved very much
further and had lingered on in South Africa, to be eventually ab-
sorbed by the oncoming ancestors of the living Bushmen.
The Border Cave Skull 1
I n 1934 Raymond Dart dug a trench in a cave on the border be-
tween Swaziland and Zululand (because it is on the border it is
called the Border Cave), and in 1940 W. E. Horton of Nsoko,
Swaziland, while continuing the excavation, found a human fron-
tal bone in a late Stillbay cultural context. Further excavations
and screening in 1941 yielded additional pieces of the skull, which
is still fragmentary, consisting mostly of a frontal bone and parts
of a left parietal, temporal, and occipital. Its basic measurements
are given in Table 37.
The Border Cave skull is 200 cc. larger than the Cape Flats skull
and has a vault 8 mm. higher. Its brow ridges are still large, and
its general morphology is as expected if the Chellian-3 to Rhode-
sian line continued to evolve past the Cape Flats grade. It looks
no more like a Bushman than did the late Mrs. Jones, with whose
remark we began this chapter.
Writing before the discovery of the Chellian-3 skull, Briggs saw
in the Border Cave cranial vault a resemblance to the Mouillians
1 H. B. Cooke, B. D. Malan, and L. H. Wells: “Fossil Man in the Lembobo
Mountains, South Africa; the Border Cave, Ingwavuma District, Zululand,” Man,
Vol. 45, Art. No. 3 (1945), pp. 6-12.
Wells: “Photographs with Note: The Border Cave Skull,” AJPA, Vol. 8, No. 2
(i95°)> PP- 241-3.
The Border Cave Skull
633
of Afalou-bou-Rhummel.2 His interpretation does not conflict with
mine, however, because in the part of the skull represented by the
Border Cave specimen Caucasoids (including the Mouillians)
and Congoids have much in common. Ronald Singer, the dis-
coverer of the Saldanha Bay skull, permits me to say that he is not
convinced that either the Cape Flats or the Border Cave skull be-
longs to any group ancestral to the Bushman.3
There can be little doubt that the Cape Flats and Border Cave
skulls were those of Negroes, and the Cape Flats skull had not
evolved far beyond the erectus-sapiens threshold. We can inter-
pret the presence of these skulls in South Africa in post-Pleisto-
cene time in two ways. Either they represent an invasion or infil-
tration of Negroes from the north, along with the complex of cat-
tle-breeding and metal-using which gave rise to the culture of the
Hottentots; or else they were the remnants of a local population
that had evolved from the Chellian 3-Saldanha-Broken Hill line
into a local race of Negroes, one which became extinct by absorp-
tion into the ranks of the invaders.4
Whether or not a local race of Negroes evolved in South Africa
before the ancestors of the Bushmen arrived has little to do with
the origin of the Congoids as a subspecies. South Africa was not
the principal center of the evolution of that or any other subspe-
cies. The Negro home has traditionally been West Africa, a part
of the Dark Continent from which we have not a single scrap of
evidence.
2 Briggs: “The Stone Age Races of North Africa,” p. 65.
3 Personal communication, January 18, 1962.
4 To this second interpretation may be added a piece of evidence worthy only
of a footnote. In or about 1941 a local naturalist named W. E. Jones excavated a
nearly complete skeleton in a gorge of the Tugela River in Natal. It was buried
in warm moist acid soil of a type inimical to the preservation of bone for more
than a few centuries. Furthermore, the burial was accompanied by ostrich eggshell
beads of Bushman type, which would not have lasted long in that soil. According
to Jones, the skull was very similar to the Broken Hill specimen, particularly in
the size of its brow ridges and the slope of its forehead. Realizing the importance
of his discovery, Jones gave the skull to a Dr. Warren, director of the Natal Pro-
vincial Museum, who apparently took it to London during the latter part of
World War II. Dr. Warren died in London and the skull has disappeared.
O. Davies: “A Missing Skull of Early Type from Zululand,” Man, Vol. 57, Article
54 (1957), P- 48.
634
Africa
The Capsian Settlers of the White Highlands 5
In the highlands of Kenya, Tanganyika, and Ruanda-Urundi,
where the altitude is from 5,000 to 7,000 feet, the weather is cool
the year around. Early in the present century a number of Euro-
peans set up farms in the coolest parts of this region, and built
Nairobi. Thousands of years earlier this same climate seems to
have attracted other Caucasoid immigrants, a tall, long-faced,
narrow-nosed people who buried their dead in a contracted pos-
ture and made blade tools in the Capsian style. We have seen
these people and their tools before, in North Africa.
The first of these burials was discovered by Hans Reck of Berlin
in Olduvai Gorge, in soil that Leakey has since identified as Bed V
( see Fig. 40), a Capsian level. The second site was a series of buri-
als along the face of a cliff on a farm in the Elmenteita district of
Kenya. They were discovered in 1918. After the bodies had been
buried, the waters of Lake Nakuru had risen, disturbing and re-
depositing the skeletons, and then later had subsided. At least
thirty individuals were interred in this ancient cemetery.
In Kenya Leakey found the third site, Gamble’s Cave, after
which the Gamblian Pluvial period was named. He excavated five
skeletons from this cave. Mrs. Leakey and A. J. Poppy discovered
the fourth site, which contained a skeleton of the same type as the
others, lying in the edge of an ancient lake at Naivasha Railroad
Station, Kenya. Except for Olduvai, these sites are close to each
other ( see Map 12 ) .
Only Gamble’s Cave is completely stratified, and it alone con-
tained adequate samples of implements and fauna. The imple-
ments are Capsian throughout, but between the second and fourth
occupation levels from the top, an intrusive stratum of African
Middle Stone Age tools was found, with Capsian artifacts below
and above it. The fauna was modern.
5W. Giesler and T. Mollison: “Untersuchungen iiber den Oldowayfund,”
VGPA, Vol. 3 ( 1929), pp. 50-67.
Leakey: The Stone Age Races of Kenya.
Leakey: “The Naivasha Fossil Skull and Skeleton,” JEAN, Vol. 16, No. 4-5
(1942), pp. 169-77.
S. Cole: The Prehistory of East Africa (London: Pelican Books; 1954).
Briggs: “The Stone Age Races of Northwest Africa.”
McBumey: The Stone Age of Northern Africa.
The Capsian Settlers of the White Highlands 635
The key to the age of this site lies in three facts. Potsherds were
found in two of the three Capsian layers. Pottery was not made in
the Near East or anywhere else, except Japan, before 5,400 b.c. As
the Capsians were a culturally peripheral folk, it is unlikely that
they were the first to invent pottery. Also, in an upper layer was
found a bone harpoon identical with others from an upper layer
in a Congo site dated at 6,000-4,500 b.c.6 All the other Capsian
sites in Kenya are postglacial. Therefore, the Gamble’s Cave site
fits into the local time scale where it belongs, well after the arrival
of the Capsians in North Africa.
How the Capsians got to East Africa from North Africa we do
not know, but we do not really need to. There is a direct overland
route across the Sahara, following a diagonal path across the Ti-
besti Highlands. There is also the Nile Valley. At the time of their
migration, during the early millennia of the post-Pleistocene, the
Sahara had more surface water than it has today and it supported
herds of game and was more densely populated than it has been in
recent times.
Eleven skulls from the four sites are all essentially Caucasoid
(see Table 37). They belonged to a rugged form of the Medi-
terranean race (a division of the Caucasoids), with long brain
cases, narrow faces, and long noses. They all have modern-style
chins, and in the males the mandibles are deep and the gonial an-
gles everted. Most of them could pass unnoticed in a collection
of Capsian skulls from North Africa. Between the males and fe-
males, considerable differences are evident. This is best indicated
in their respective cranial capacities, which average 1,497 cc. f°r
seven males and 1,223 cc- f°r three females.
Although their teeth have not been systematically studied,
we know that no incisors were shoveled and that one milk molar in
the Elmenteita series has a Carabelli’s cusp. A lower first molar of
an Elmenteita male has the extraordinarily large crown dimen-
sions of 13 by 13 mm., well above the Caucasoid range. Also, the
Elmenteita skulls, which Leakey considers younger than those
from Gamble’s Cave, are more prognathous than the others. To
6 J. de Heinzelin: “Les Fouilles de Ishango,” Exploration du Parc National Al-
bert (Brussels: Institut des Parcs Nationaux du Congo Beige; 1957), pp. 64-1.
K. P. Oakley: “Bone Harpoons from Gamble’s Cave, Kenya,” The Antiquaries
Journal, Vol. 41, Nos. 1-2 (1961), pp. 86-7.
636 Africa
my eye it looks as if they were the product of a mixture between
the invading Capsians and an earlier element, either ancestral
Bushmen who had preceded them across the Sahara or people
like those whose skulls we have already seen in Cape Flats and
the Border Cave.
These people were taller than the North African Capsians. The
Olduvai V skeleton was that of a man five feet eleven inches tall
(180 cm.); No. 4 from Gamble’s Cave was the same or a trifle
taller; and the Naivasha Bailroad Station man, according to
Leakey, was six feet eight inches tall (203 cm.)— a giant. These
statures compare well with those of the living Watusi and other
tall Hamitic peoples of the present-day white highlands.
Leakey and his associates also excavated a number of Neolithic
sites in the same region. At least some of them were agricultural,
and all antedate the main push of the Bantu expansion. These
sites include the Nakuru burial ground, with nine skeletons, one
of which was measured, and Willey’s Kopje, near Elmenteita, with
three skeletons, all measured. All twelve indicate that the Cauca-
soid racial type of the earlier Capsian invaders persisted in the
highlands until the Iron Age, or until the Bantu arrived. What-
ever mixture took place was probably with the old, indigenous
East and South African line and with ancestral Bushmen rather
than with modern Negroes. Despite the predominance of Bantu
speech and culture in the highlands today, many of the native
tribesmen who have black or brown skins and Negroid hair still re-
tain Caucasoid facial features.
The Origin of the Capoids 7
The Capoids, named by Broom after the Cape of Good
Hope, constitute one of the five subspecies of modern man as
7 Dart: “Three Strandloopers from the Kaokoveld Coast,” SAJS, Vol. 51, No. 6
(i955), PP- 175-9-
Drennan: “The Dentition of a Bushman Tribe,” AS AM, Vol. 24 (1929) pp
61-87.
H. S. Gear: “A Further Report on the Boskopoid Remains from Zitzikama,”
SAJS, Vol. 23 ( 1926), pp. 923-34.
Gear: “Cranial Form in the Native Races of South Africa,” SAJS, Vol. 26
(1929), pp. 684-97.
The Origin of the Capoids 637
stipulated in Chapter 1. They include the living Bushmen, the
living Hottentots and that branch of the Hottentots known as the
Korana, a few beachcombing remnants of an earlier coastal Bush-
man population known as Strandloopers, and certain relict popu-
lations in Tanganyika and possibly farther north.
Capoid skeletal material is abundant in South Africa. Among it
may be found specimens similar to contemporary Bushmen, with
their reduced stature, small brain bent at the base to produce a
bulging forehead, and small, neotenously infantile, very flat face.
In addition there are specimens of a larger and less infantile va-
riety commonly known as the Boskop race. To these two types
may be added individual skulls and groups of skulls which show
mixture with some other element, Hamitic (like the highland
skulls we have just studied) or Negro or “Australoid.” The result
is a considerable confusion.
Two principal theories have been advanced to explain the ori-
gin of the Capoids. The first is that their ancestors came from
North Africa or the Sahara, mostly if not entirely in postglacial
A. J. H. Goodwin: “A Comparison Between the Capsian and South African
Stone Cultures,” ASAM, Vol. 24, Part 1 (1929), pp. 17-32.
Keith: New Discoveries Relating to the Antiquity of Man.
Keith: “A Descriptive Account of the Human Skulls from the Matjes River
Cave, Cape Province,” TRSS, Vol. 21 (1933), pp. 151-85.
Leakey: The Stone Age Races of Kenya, 1935.
J. F. Schofield: “The Age of the Rock Paintings of South Africa,” SAAB, Vol.
3, No. 12 (1948), pp- 79-88.
G. W. Grabham: “Note on the Geology of the Singa District of the Blue Nile,”
Antiquity, Vol. 12, No. 46 (1938), pp. 193-5.
Singer: “The Boskop ‘Race’ Problem,” Man, Vol. 58 (1958), pp. 1-5.
D. Slombe: “The Osteology of a Bushman Tribe,” ASAM, Vol. 24 (1929),
PP- 33-6o.
G. Eliot Smith: “The Influence of Racial Admixture in Ancient Egypt,” ER,
Vol. 7, No. 3 ( 1915), pp. 163-83.
P. V. Tobias: “Bushmen of the Kalahari,” MAN, Vol. 57 (1957), pp. 33-40.
L. H. Wells: “The Status of the Bushman as Revealed by a Study of Endo-
cranial Casts,” SAJS, Vol. 34 (1932), pp. 47-58.
Wells: “The Fossil Human Skull from Singa,” FMA, No. 2 (1951), pp. 29-42.
A. S. Woodward: “A Fossil Skull of an Ancestral Bushman from the Anglo-
Egyptian Sudan,” Antiquity, Vol. 12, No. 46 ( 1938), pp. 190-3.
R. Biassutti: “Crania Aegyptica,” AAE, Vol. 36, Fasc. 2 (1906), pp. 165-73.
V. Giuffrida-Ruggeri : “I Crani Egiziani del Museo Civico di Milano,” AAE,
Vol. 37 ( 1907), pp. 399-410.
F. C. Shrubsall: “Notes on Some Bushman Crania and Bones from the S.
African Museum, Capetown,” ASAM, Vol. 5, Part 5, No. 6 (1906-10), pp. 227—
70.
638
Africa
times, pushed south by the expanding Capsians. The second is
that they evolved from local ancestors, including not only Sal-
danha Bay man and Broken Hill man, but also Florisbad, a fossil
skull which will be described presently. This evolution consisted
of two steps. First they acquired large, very modern brain cases,
and then they shrank, through infantilism partly accompanied by
dwarfing. At the same time, according to this second theory, these
indigenous ancestors mixed sporadically with Hamites, “Austra-
loids,” and possibly others who wandered into the marginal terri-
tory that some of the Bushmen still occupy.
The proponents of both theories admit that the Bushmen are
descended from full-sized ancestors, and that they began to grow
small and infantile only a few thousand years ago, and that they
did so in South Africa. No one has yet offered a satisfactory ex-
planation of why they shrank. This subject, discussed in Chapter
3 (see pp. 112-15) and later in this chapter, does not concern us
here. Our present task is to trace the history of the skeletons of
Bushmen, large and small, in time and space.
The first theory, that of a northern origin, is based on data con-
cerning chronology, archaeology, and the geographical distribu-
tion of Capoids, dead and alive. First, in all of South Africa there
is not a single Bushman-like skull or skeleton of geological antiq-
uity, although a few, such as Boskop, cannot be dated one way or
the other. Second, as Goodwin has shown, all the Bushman-like
specimens that are associated with a stone-tool industry are ac-
companied by artifacts known to South African archaeologists as
Wilton, and Wilton is nothing but a derivative of the Capsian of
North and East Africa. Third, the famous Bushman rock paint-
ings, which extend northward into Bhodesia, cannot be much
more than a few centuries old because they do not extend below
the soil level in the caves and rock shelters in which they are
found; because they have not been smoke-blackened by herds-
men’s fires; because the granite on which they were painted had
not exfoliated (peeled off) very much from weathering; and be-
cause the fauna depicted is all recent.8
The theory that the ancestral Capoids migrated southward
from North Africa goes back to the discovery, during the last cen-
8 Schofield: “The Age of the Rock Paintings. . . .”
The Singa Skull from the Sudan 639
tury, of Bushman-like rock paintings in the Sahara. In 1905 Bias-
sutti first noticed Bushman-like traits in some of the oldest ancient
Egyptian skulls, and since then the theory has been proposed and
rejected several times. This is the first time, I believe, that the evi-
dence favoring it has been presented as a unit.
The Singa Skull from the Sudan
I n 1924 the theory of a northern origin was re-enforced by the
discovery, made by W. R. G. Bond, of a Bushman-like skull at
Singa, 200 miles south of Khartoum, on a bank of the Blue Nile.
Grabham, who had studied the site geologically from the stand-
point of rates of soil deposition caused by the overflow of the Nile,
stated, in 1938, that it could not be less than 5,000 nor more than
10,000 years old. The skull, however, was completely mineralized,
despite the fact that it is thought to be of fairly recent date.
In 1951 Miss Dorothea Bate,9 who studied the animal bones,
found that two and possibly three supposedly extinct species of
mammals were contemporaneous with the skull. They are Homoi-
oceras singae Bate, an extinct antelopine; Hystrix atasohae Bate,
an extinct porcupine; and Sivatherium , an extinct member of the
giraffe family. Because no other specimens of the first two species
had ever been found, we cannot be sure when they became ex-
tinct, and she herself was not altogether certain that she had
correctly identified Sivatherium, which did not survive the
Pleistocene.
In any event, the Singa skullcap is clearly as old as, if not
older than, any known and competently dated Capoid skull
found yet in South Africa. The skullcap is nearly complete, but
the face is missing. The bone is thick (13 mm. on the parietals)
and the brow ridges moderately heavy, with a distinct notch
above glabella, like Sinanthropus and like the least infantile of the
Bushman skulls. The forehead is narrow, but bulging; the parie-
tals also bulge, giving the brain case a pentagonoid appearance.
The orbits were apparently rectangular. Morphologically the skull
could have been that of a full-sized progenitor of a Bushman. It is
9 Wells: “The Fossil Human Skull. . .
♦
Africa
640
brachycranial ( length = 188; breadth = 154; cranial index =
82) but it may have been distorted. Even if the index of 82 is cor-
rect, this does not invalidate the racial identification of the Singa
skull, because there are some living brachycephalic Bushmen. In-
deed, some of the Mouillian skulls from Algeria and Morocco are
also voluminous, brachycranial and rugged, presumably from
mixture with the local Aterian population which they partly ab-
sorbed and partly displaced.
The Homa Shell-Mound Skulls
Following the hypothetical migration route of the ances-
tors of the Bushmen to the south, we come to the Homa shell
mound, a Mesolithic midden of fresh-water mollusk shells situ-
ated on the shore of Lake Victoria in Kenya. This midden con-
tained Wilton A (Early Wilton) tools, typical of Bushmen, and
six skeletons in various states of preservation. Two of the skulls,
Numbers 1 and 4, have been measured and illustrations of them
have been published 1 ( see Table 37 ) .
Number 1 belonged to a middle-aged male who had a large
brain case, weak brow ridges, and a strongly arched forehead. Al-
though his mandible was large, his teeth were small, especially his
canines and incisors. His largest lower molar was the first one,
and the third was greatly reduced. Number 2, which consists of
postcranial bones and a mandible, belonged to a tall, massive man
with a big mandible and five cusps on all his lower molars. Num-
ber 3 is said to be a virtually complete skeleton of a short, thick-
set adult male with a large head.
Number 4’s face is nearly complete, and we have a profile draw-
ing of the whole skull. The cranial capacity is very great, and the
skull long and high. The forehead is bulbous, the top of the head
comparatively flat. The face has much alveolar prognathism, the
nasal skeleton is flat, and the lower rim of the orbits project. De-
spite its great size, this skull is morphologically that of a Bush-
man. Number 4’s body was correspondingly large and heavy.
These Homa Shell Mound skeletons are significant not only be-
cause they place the Bushmen’s ancestors well north of the mod-
1 Leaky: The Stone Age Races of Kenya.
The Boskop Brain Case and the “Boskop Race ” 641
ern range of the Capoids, but also because, along with Singa, they
are still large, unreduced, and not infantile. Had the Bushmen
spread north to Kenya after they had completed their peculiar
pattern of evolution in South Africa, as Tobias (1957) has sug-
gested, Leakey would have found small Bushmen in the Homa
mounds instead of big ones.
The Boskop Brain Case and the “Boskop Race ”
I n 1913, when the study of human paleontology was in its in-
fancy, F. W. Fitzsimmons of the Port Elizabeth Museum found
part of a brain case, a loose temporal bone, a piece of mandible,
and several bits and pieces of long bones in a layer of laterite
four feet from the surface on the east bank of the Mooi River, in
the Potchefstroom District, southwest Transvaal. These remnants
of an ancient burial were to become famous as the Boskop man,
which has gone into the literature as the type specimen of the so-
called Boskop race.
The bones were mineralized as one might expect after they had
lain for some time in laterite. No fauna was found with them, nor
any implements, although later a single blade tool was picked up
nearby.2 The only blade industry in that region was Wilton. Al-
though the find cannot be dated, it probably belonged to the early
Capoid group which arrived in South Africa several thousand
years b.c.
As Table 37 shows, the Boskop calvarium would fit in the
Homa shell-mound population without difficulty; as it was a little
broader than Homa 1 or 2, it deviated from them in the direction
of Singa. Morphologically it possessed the earmarks of the Bush-
man in vault form and forehead shape, small mastoids, and other
details; and, like the Homa skulls, its mandible has a chin. It has
no face, but neither does the type specimen of the Neanderthals.
Anyone who wishes to call the unreduced ancestors of the Bush-
man the ‘ Boskop race” is, of course, free to do so. But I consider
the continued use of this name unnecessary.
2C. van R. Lowe: “An Artifact Recovered with the Boskop Calvaria,” SAAB,
Vol. 9, No. 36 (1954), pp. 135-7.
642
Africa
The Florisbad Cranial Fragment 3
T o those familiar with the conventional classifications of geologi-
cally ancient skulls it may come as a surprise to see the Florisbad
cranial fragment treated separately from the Saldanha and
Broken Hill skulls, with which it is generally grouped in a fossil
triumvirate. Actually, Florisbad has little in common with the
other two skulls except birth in Africa and membership in the ge-
nus Flomo. Two factors have created this confusion: inconclusive
dating and faulty reconstruction.
In 1932 T. F. Dreyer was excavating the complex deposits of
the site of a mineral spring named Florisbad, 25 miles north of
Bloemfontein in the Transvaal. At a depth of 20 feet he found a
fragmentary cranium consisting of most of the frontal bone, parts
of the parietal and occipital, various pieces of the facial bones,
mostly from the right side, and a loose right third upper molar.
The position of the skull in the site was unusual (see Fig. 83)
because of the nature of the spring. The spring consists of a row
of eyes or fountains that have taken turns flowing to the surface at
different times. Periodically the active eye stops flowing and a
layer of peat forms over it. At the eye where the skull was found
there were four layers of peat, numbered I to IV, starting at the
bottom. Layers II, III, and IV had been deposited after the eye
had died, but layer I was older than the eye, which had broken
3 T. F. Dreyer: “A Human Skull from Florisbad,” PASA, Vol. 38 (1935), pp.
119-28.
Dreyer: “Endocranial Cast of the Florisbad Skull,” SNNM, Vol. 1 (1936),
pp. 21-3.
Dreyer: “The Fissuration of the Frontal Endocranial Cast of the Florisbad
Skull Compared with that of the Rhodesian Skull,” ZfRK, Vol. 8 (1938), pp.
129-98.
Drennan: “The Florisbad Skull and Brain Cast,” TRSS, Vol. 25 (1937), pp.
103-14.
A. Galloway: “Man in Africa in the Light of Recent Discoveries,” SA/S, Vol.
34 (1937), PP- 89-120.
Galloway: “Nature and Status of the Florisbad Skull as Revealed by its Non-
metrical Features,” AJPA, Vol. 23, No. 1 ( 1937), pp. 1-16.
Oakley: “New Evidence Regarding Rhodesian (Broken Hill) Man.”
Oakley: “The Dating of the Broken Hill, Florisbad, and Saldanha Skulls.”
Oakley: “Dating the Stages of Hominid Evolution.”
Singer: “Man and Mammals in South Africa.”
Singer: “The Rhodesian, Florisbad, and Saldanha Skulls.”
The Florishad Cranial Fragment
643
through it from below, bringing with it sand, animal bones, and
probably also artifacts. The skull lay in a depression in the top of
the first layer, next to the edge of the eye hole. Resting on the
top of the peat of layer I was a perfectly modern-looking grind-
stone and muller ( metate and mano to American archaeologists ) .
The most logical interpretation of this unusual situation is that
the cranial fragment, which judging by toothmarks had been part
SANDY SOIL
PEAT IV-
\Ubjjfo M \ M..< .4lf,
— ■ : :V~. r v:Afr Vi‘V'» ■■■ /
YELLOW SAND-
mmm
GREENISH SAND-
QUERN-
PEAT I
BROWN SANDV |
BASAL PEAT-
OLD EYE OF SPRING
FOSSILS + BASALT TOOLS
Fig. 83 The Florisbad Site. The Florisbad skull site is a thermal spring with
many eyes, most of which ceased to bubble thousands of years ago. The Florisbad
skull fell into one of these eyes while it was still active. Later the flow of mineral
water ceased, and three sets of alternate layers of sand and peat were laid down
above it. Because the skull is younger than Peat I, which is under it, and the
overlying undisturbed Peat II, its age depends on the dates given the two layers of
peat. ( Drawing after Oakley, 1954.)
of a carnivore’s meal, had been dropped into the spring after the
water had burst through layer I and before the formation of
layer II. It thus is not necessarily associated with any of the ani-
mal bones or artifacts, except for the grindstone and muller, which
could hardly have been cast up by the spring. The dates of these
peat layers are therefore critical.
In 1954 Libby published the following Carbon- 14 dates:
Layer III = 6,700 ± 500 years (C-852); Layer II = 9,104 ± 420
644 Africa
years (C-851); Layer I = older than 41,000 years (C-850). In
195® the Lamont Laboratory of Columbia University published
the following: Layer III = 19,530 ± 650 years (L-271D); Layer
II — 28,450 ± 200 years (L-271C); Layer 1 = 35,000 years
(L-271B). Libby tested solid carbon; the Lamont Laboratory
used carbon dioxide. Which one, if either, of these sets of figures
is right?
In 1955> E. M. van Zindern Bakker, a paleobotanist who worked
on the Florisbad peats without reaching a definite conclusion, had
this to say about using the Carbon- 14 method on the Florisbad
peat : The production of much methane by the spring shows that
the water in percolating through the rock must have contact with
Ecca coal deposits or with carbonaceous shales or their distillation
products formed under the influence of the intruding dolerite.
This contact may have had an important influence on the result of
the age determination, as indicated by Oakley. The dark layers
may be dated too old as they may contain ‘old carbon’ from the
above-mentioned Ecca strata.”4 His pollen analysis indicated
that the peat of layer I was probably Upper Pleistocene and that
much time elapsed before layer II was formed, during the post-
Pleistocene Makalian wet period, the very time in which the peo-
ple of Gamble’s Cave lived.
In short, the Florisbad skull could have been dropped down the
eye of the spring as late as 7,000 to 5,000 b.c. This terminal date
would be consistent with the anatomical identification of Floris-
bad as an ancestral Bushman, which it seems to have been.
Dreyer reconstructed the fragment by adding plaster to the
original bone, of which no cast had been made. Probably with a
Neanderthal image in mind, he tilted the bone backward and
stretched out the maxilla, adding to the actual tooth sockets to
produce an elongated, prognathous upper face. The result was an
unnatural composite that has plagued textbook writers for three
decades.
After Dreyer ’s death, A. C. Hoffman, director of the National
Museum of Bloemfontein, had the plaster removed and six casts
were made of the original bone, one of which he presented
4E. M. van Zinderen Bakker: “A Pollen Analytical Investigation of the Floris-
bad Deposits (South Africa),” PTPA (1957), pp. 56-67.
The Formation of the Modern Capoid Peoples 645
to me (see Plate XXXI).5 The work was done in the Depart-
ment of Anatomy of the University of Capetown. Seen in its
original form, the Florisbad fragment is part of a large sapiens
skull, with a sharply curved frontal bone; a very broad forehead; a
short, orthognathous face; square orbits; and teeth of modern size,
judging by the sockets and by the remaining upper third molar.
It would fit in the Homa series; it resembles Boskop; and it pat-
ently belonged to an ancestral Bushman. Further details may be
expected from Hoffman. If the fragment is older than I think it is,
I shall be surprised, and my reconstruction of the origin of the
Capoids will be dealt a serious blow.
The Formation of the Modern Capoid Peoples
The remainder of the skeletal material from South Africa
does not present serious problems. Owing to the diligence and en-
thusiasm of local archaeologists, many prehistoric Capoid skele-
tons have been discovered. The best known are Fish Hoek, a
complete skeleton; the collection from Zitzikama, consisting of
five imperfect specimens; and the Matjes River group, consisting
of eighteen skeletons in various states of preservation. Some of
these may be as old as Florisbad or older, but all are without much
doubt post-Pleistocene.
This material indicates that the process variously known as
shrinking, fetalization, or general size reduction, which affected
the Capoids, took place in large part, if not completely, after their
arrival in South Africa. Fish Hoek has a large brain case, but his
face is small. One Zitzikama specimen, possibly hydrocephalic,
has a very bulging forehead and a tiny face. In the Matjes River
group there are full-sized and shrunken individuals in the same
population. Singer ( 1958 ) has pointed out that even among liv-
ing Bushmen and Hottentots individuals who fit the ancestral ra-
cial dimensions may be found.
The size reduction began no more than eight or nine thousand
years ago and has not yet affected the entire Capoid population to
5 Hoffman has invited five physical anthropologists, including myself, to make
our own reconstructions while he makes a sixth. He plans to compare the results.
Africa
646
the same degree. The coastal Bushmen, or Strandloopers, a few
of whom survive in southwest Africa and Angola, were taller and
had larger brains than the Bushmen of the Cape and Kalahari.
Moreover, as one moves northward in South Africa and Bechua-
naland, the Bushmen grow taller. In South Africa itself the Hot-
tentots are less infantile and a little larger than the Bushmen.
Why this is so, I am not prepared to say, but two possibilities sug-
gest themselves. The Hamitic racial element which the Hottentots
are believed to have acquired along with their cattle may have
Fig. 84 The Capoid Line: Profiles of Homa, Fish Hoek, and a Modern
Bushman. A. Skull 4 from the Homa Shell Mounds in Kenya (after Leakey, 1935);
B. Fish Hoek (after Drennan, 1929, and a cast); C. A Bushman, the so-called
“Hottentot Venus” (after de Quatrefages and Hamy, 1882). According to the
theory presented in this book, the Capoids originated in North Africa as the
Temefine-Tangier line, and moved to East and finally South Africa when pushed
out by the invading Mouillians and Capsians, who were Caucasoid. Capoid skulls
have been found all the way from the Sudan ( Singa ) to the Cape of Good Hope.
At first large and rugged, they gradually become small and infantile. Yet they
preserved one special facial feature — facial flatness, which may be observed in the
three specimens shown here.
slowed down the process, or the pastoral life may in some way fa-
vor small size and infantilism less than hunting does.
Frankly, I am unable to explain the pedomorphism and partial
dwarfing of the Bushmen. They do not live in tropical forests and
apparently never did. It can have nothing to do with food
economy because before they were disturbed in South Africa
they had all the game they could eat. A parallel might be drawn
between them and the Lapps, who are also stunted and tend to
have small faces and small teeth. The Lapps occupy the most
poleward part of their racial realm; the Bushmen the most pole-
The Formation of the Modern Capoid Peoples 647
ward of theirs. Under certain circumstances there must be some
selective advantage in pedomorphy, or it would not have become
general in either of these unrelated populations.6
Pedomorphy did not spread among the Capoid peoples who
lived north of its South African center of origin, as we know from
excavations at two modern, Metal Age sites in northern Transvaal
near the Limpopo River. These sites are Mapungubwe and Bam-
bandyanolo, two neighboring hilltop forts or shrines excavated be-
tween 1933 and 1940.7 These sites were occupied between a.d.
1,000 and 1,400, as established by Carbon-14 tests.8 Their inhabit-
ants were Metal Age people, undoubtedly food-producing, who
wore gold ornaments. By a.d. 1,400 the younger of the two sites,
Mapungubwe, is known to have been inhabited by Bantus, and
the local Bantus still hold these hills in reverence.
Galloway has studied seventy-four skeletons from Bambandy-
anolo and eleven from Mapungubwe. Not one was that of a Ne-
gro. All were full-sized Capoids, some of whom had taurodont
teeth. Clay figurines found with the skeletons show the elongated
labia minora and steatopygia characteristic of Bushmen and Hot-
tentots. Galloway’s careful work on these skeletons also illustrates
the differences between Capoids and Negroes in many details of
anatomy, particularly in respect to the vertebrae and the bones of
the feet. These show that the two subspecies are no more alike in
the postcranial skeleton than in the skull, or in the anatomy of the
soft parts.
The skeletons which Galloway described indicate that an un-
reduced Capoid population lived on the banks of the Limpopo
6 A fantastic idea that I give only in a footnote stems from the fact that the
Bushmen use poison. Every man normally has enough of it on his person to kill a
rival in a few minutes during the victim’s sleep. This knowledge, according to
Mrs. Lorna Marshall, who has lived among them, is a powerful deterrent to
adultery. Whether or not the self-control needed to refrain from adultery, and
therefore avoid being killed, has an endocrine basis which favors pedomorphy I
do not know, but I do know that adultery is said to be rare among Andamanese
and other Pygmies.
7 Galloway: “The Skeletal Remains of Mapungubwe,” in L. Fouche, ed:
Mapungubwe (Cambridge: Cambridge University Press; 1937), pp. 127-74.
Galloway: “The Skeletal Remains of Bambandyanalo” (Johannesburg: Wit-
watersrand University Press; i960).
8 Mapungubwe = a.d. 1430*60 (Y-135-9) and a.d. 1390*60 (Y-135-14);
Bambandyanolo = a.d. 1060*65 (Y-135-17).
Africa
648
only two hundred years before the Dutch settled Capetown, and
if these skeletons were those of Bantu-speakers, then the early
waves of Bantus across the Limpopo must have been thin indeed,
like the first waves of Mongoloids into southeast Asia.
It should be noted also that several populations between Libya
and the Rhodesias still show traces of Capoid origin or at least
Capoid mixture. The first of these is a small group of desert dwell-
ers called Duwwud, or Worm-People in Arabic, who live on the
banks of three salt lakes in the Fezzan, Libya.0 The Duwwuds,
black-skinned, curly-haired people, specialize in netting Artemia,
the brine shrimp, in vast numbers in September, when these ani-
mals reach their seasonal peak of abundance. The women pound
the catch into a shrimp paste which they pat into cakes and let
dry. These cakes are sold to Arab caravan men. During the rest of
the year the Duwwuds run about the dunes hunting jerboas and
other small game, which they kill with thrown sticks. In Cipriani’s
photographs, some of these people resemble the classic descrip-
tion of a Boskop type, with a flat face and a big jaw. Others show
traces of Arab, or Negro, admixture. Although this is not concrete
evidence of the former presence of an unreduced Capoid popula-
tion in Libya, it is at least a lead that would warrant further inves-
tigation. According to Briggs, the Sahara contains more than one
such population.1
Better evidence is afforded by the presence in central Tangan-
yika of a tribe that still speaks a Bushman-Hottentot language.
They are the Sandawe, some of whom were measured in 1944 by
J. C. Trevor of Cambridge University.2 Although somewhat mixed
with Bantu, the Sandawe still resemble the Capoid peoples, and
particularly the Hottentots, anthropometrically. From the zoo-
geographical standpoint it is much easier to interpret the Sandawe
living today in central Tanganyika as a relict population left be-
hind during a southward movement than as the result of a north-
ward push against the main current of migrations in East Africa,
9 L. Cipriani: “Un Interesante Pueblo del Sahara: los Dauada,” RGA, Vol. 2,
No. 2 (1934), PP- 141-52-
1 Briggs: Tribes of the Sahara (Cambridge, Mass.: Harvard University Press;
i960).
2 J. C. Trevor: “The Physical Characters of the Sandawe,” JRAI, Vol. 77, Pt. 1
(i947)» PP- 61-80.
The Earliest Skeletons of Modern Negroes 649
particularly since Bushman rock paintings, perforated stones, and
so on, are not found that far north.
All the evidence that we have reviewed, then, including that of
archaeology, of skeletons, and of relict populations, would indi-
cate that the Bushmen, Hottentots, and their larger ancestors are
descended from the Ternefine-Tangier line of North Africa; that
their ancestors were driven out of that region by an invasion of
Caucasoids toward the end of the Pleistocene and during the
early post-Pleistocene period; and that they did not begin to un-
dergo a reduction in size until after arriving in their historic home-
land, South Africa, Southwest Africa, Bechuanaland, and the
Rhodesias.
The Earliest Skeletons of Modern Negroes
Now that the racial history of the Capoids has been traced
more or less satisfactorily, only one subspecies remains to be ac-
counted for, and that is the Congoid. It is the weakest warp in our
racial fabric. The ancient line running from Chellian-3 man to
to Cape Flats and the Border Cave is Negro more than anything
else, but it is not fully Negro in the modern sense. The Negroes as
we know them are a distinctive people, anatomically and physio-
logically, and must have arisen in another part of Africa, probably
north and west of the Congo basin. There evidence is scarce and
of late date.
The oldest skeleton that all writers agree was that of a Negro is
the so-called Asselar man, found in 1927 by M. V. Besnard and
T. Monod in the dry bed of a once wide and perennially flowing
river near the Tilemsi depression in the Sahara, 400 kilometers
north of Timbuktu at 190 N. latitude and a little east of the Green-
wich Meridian, latitude o° 3
No implements were associated with this skeleton. With it,
however, were found the remains of fresh-water molluscs, fish,
crocodiles, and various gazelles and antelopes, all of which still
exist, but not in the Sahara. It obviously dates back to a time
when this part of the Sahara was well watered, and is probably
3 Boule and Vallois: “L'Homme Fossile d’ Asselar, Sahara,” A1PH, Mem. No. 9,
1932-
Africa
650
post-Pleistocene. It was that of an adult male at least five feet
seven inches tall ( 170 cm. ), whose long bones were slender, whose
forearms were long in relation to his upper arms, and whose lower
legs were long in relation to his thighs. His pelvis, vertebrae, and
hand and foot bones were all Negroid. In fact, from the neck
down he was altogether a Negro.
His skull is not so easily categorized. Its capacity of 1,520 cc.
lies on the high side of the Negro range, and its vault proportions
can be matched today in Negro skulls from the Sudan, particularly
in a series of Wolof skulls with which Boule and Vallois compared
the skull of Asselar. As for the face, it deviates from that of the
Wolof s only in its shortness, but short faces are common in West
Africa. Its two upper median incisors had been removed soon
after eruption, presumably in some initiation rite. This loss caused
considerable bone damage and may have reduced the upper face
height ( nasion-alveon ) .
The teeth of Asselar man are typically Negro in dimensions and,
as far as we can tell, in form. In the upper jaw the second is the
largest molar, followed in size by the first and then the third. In
the lower jaw the ranking is second, third, and first, which is
both primitive and unusual. In the upper jaw only the third is
preserved well enough to permit the counting of cusps. It has
four. In the lower jaw all three molars have five cusps each.
In this skeleton Boule and Vallois saw a type of Negro closer to
the South African Bantu of today than to the living Negroes of
the Sahara and western Sudan. They attributed this postulated
relationship to mixture between a true Negro and an ancestral
Bushman before the latter had journeyed southward. They may
be right; but to me Asselar is simply a Negro, the first true Negro
that we have found in Africa who can meet all the specifications
of his race.
Remains of at least ten other individuals associated with Neo-
lithic cultural materials, and possibly Mesolithic materials as well,
have been found in a rock shelter at Kourounkorokale in Mali, in
a part of what used to be the French Sudan, 37 kilometers south-
west of Bamako in the Mandingo Hills.4 Although these speci-
4 G. Szumowski: “Fouilles de I’Abri Sous Roche de Kourounkorokale (Soudan
Frangais), BIAF, Vol. 18, Ser. B., No. 2-3 (1956), pp. 462-508.
Do the Pygmies Hold the Answer? 651
mens have not yet been studied, preliminary observations indi-
cate that they are Congoid and that some are of very small stature.
A specimen stated to have been that of a Negro was found in
1948 in Khartum, near the railroad station. Several skeletons had
been buried in a mound along with Mesolithic stone tools and
with pottery. According to McBurney, the Khartum Neolithic be-
gan about 3,253 ± 295 b.c.,5 and the Mesolithic material, includ-
ing the pottery, could hardly be much more than 500 years
older.6 One skull has been partly described.7 It is Negroid, but to
me it looks like the skull of a modern, local Sudanese, a mixture
of Hamite and Negro, rather than like the skull of a full Negro. As
these burials may not have been much more than a thousand years
older than the earliest Egyptian mural representations of Negroes,
the presence of a Negro or Negroid in the Sudan at 3,700 to
4,000 b.c. is not surprising.
Do the Pygmies Hold the Answer P
Until now we have neglected one of the four principal racial
assemblages in Black Africa, that of the Pygmies. They are esti-
mated to number about 168,500 persons 8 living in several isolated
portions of the tropical forests of Central Africa from Gabon
and the Cameroons to Uganda and Ruanda-Urundi, nearly all
north of the Congo River. The western, central, and northeastern
Pygmies live in the lowland forests, the southeastern in the up-
land forest around and on the slopes of Mount Ruwenzori.
There is some historical evidence to indicate that the western
Pygmies once extended along the entire west coast of Africa as
far as Liberia, and that as late as the sixteenth century the Pyg-
5 This date is apparently a combination of two dates: C-753 = 5,060 ± 450
B P- for charcoal from Shaheinab; and C-754 = 5,446 ± 380 B.P. for shell from
the same site. The average of the two is 5,253 ± 415 B.P., or 3,253 ± 295 b.c.,
if 2,000 years are subtracted in conversion from B.P. to B.C.
6 McBurney: The Stone Age of Northern Africa (New York: Pelican Books;
i960), p. 244.
7 D. E. Derry: “Report of Human Remains,” in A. J. Arkell: Early Khartoum
(London: Oxford University Press; 1949), pp. 31-3.
8 M. Gusinde: “Pygmies and Pygmoids,” AQ, Vol. 28 (NS Vol. 3), No. 1
(1955), PP- 3-6i.
Africa
652
mies were the principal if not the only inhabitants of the forest be-
tween Lakes Albert and Edward.
Because we have no Pygmy skeletal material other than the re-
mains of the recently deceased, and no archaeological industries
that can be attributed, with confidence, to the Pygmies, we know
nothing about these little people except that they have lived in the
equatorial forests of Africa for as long a time as is covered by the
records of history. Nor is there any evidence that Negroes lived in
the forests with them before the Negroes had acquired iron and
agriculture.
Since entering the forests Negro men have taken Pygmy wives
and there are whole tribes of Negroes that are part Pygmy in
origin and appearance; but the Pygmy men have not married
Negro women, and even if a Negress should bear a child to a
Pygmy the child would remain in the village and be considered a
Negro. There are no hybrids in the Pygmy camps in the deep
forests. To live like a Pygmy you have to be one. Gene flow be-
tween Pygmies and Negroes is thus a one-way stream which may
have made the Negroes biologically more adaptable to forest liv-
ing than were their ancestors out on the savannahs and grass-
lands.
The sickle-cell trait, an inherited malformation of the red cor-
puscles that limits their ability to carry oxygen and renders men
immune to malignant malaria, is found among 26 per cent of the
Pygmies living in malarial regions. Although not the highest fre-
quency in Africa, this figure suggests that the trait may have
originated among the Pygmies: owing to the unique direction of
gene flow between Pygmies and Negroes, the Pygmies could not
have got it from the Negroes.
In the Stanleyville region, including the Ituri forest, the home
of the eastern Pygmies, the Pygmy birth rate is constant or in-
creasing whereas that of the Negroes is declining. The survival of
Negroes in this area seems to depend essentially on a steady rate
of absorption of Pygmy genes. To the extent that forest Negroes
exchange genes with savannah and grassland Negroes, Pygmy
genes can be carried outside the forest to Negro populations that
have never lived in it.
It appears possible, therefore, that the modern Negro is, in gen-
Do the Pygmies Hold the Answer? 653
eral, part Pygmy, but he has only become so to a considerable ex-
tent since Negro cultivators first entered the forest to plant their
gardens, between two and three thousand years ago. Before that,
the gene flow between Pygmies and Negroes was probably limited
to peripheral contacts on the edges of the forests.
The Bantu tribes of East and South Africa may have been par-
ticularly affected because the Bantus originated in erstwhile
Pygmy country in West Africa and passed through the forests on
their migrations eastward and southward. The difference between
the old Negro skeletons that we have reviewed and the skeletons
of modern Negroes may reflect the presence and absence of Pygmy
genes.
If this exercise in historical reconstruction is true, then what are
the Pygmies? As they are a living people without prehistoric or
even historic skeletal antecedents, their description belongs in an-
other book, a book devoted to the living peoples of the world; but
a few facts about the Pygmies must be given here if we are ever to
unravel the racial history of Africa.9
The Pygmies are small, Negroid people. The mean stature of
the men is below 150 cm. in most groups, although those that live
in the chilly forests of Mount Buwenzori have a mean stature of
153 cm., as one would expect in view of Bergmann’s rule (see
Chapter 2). There is considerable sexual dimorphism, the women
being as a rule much smaller than the men.
They are not miniature dwarfs, like some of those in the Aus-
traloid quadrant of the Old World, nor are they as neotenous as
the Bushmen; the Pygmy men have genitals as large as those of
Negroes. Their manner of dwarfing verges on the achondroplastic
but does not reach the extremes of achondroplasia 1 seen in indi-
9 The literature is voluminous. A bibliography of 132 titles will be found in
R. R. Gates: “The African Pygmies,” AGMG, Vol. 7 (1958), pp. 159-218.
In the preparation of this section I have also directly consulted:
Gusinde: “Pygmies and Pygmoids.”
Gusinde: Die T widen. Pygmden und Pygmoide um Tropischen Afrika (Vienna:
Wilhelm Rraumiiller; 1956).
Twiesselmann : “Les Pygmees de 1’Afrique Centrale,” RM, No. 4 (1952), pp.
1-20.
Vallois: “New Research on the Western Negrillos,” AJPA, Vol. 26 (March
1940), pp. 449-71.
1 A type of dwarfing characterized by short, deformed extremities, as in bull-
dogs.
Africa
654
vidual mutations in European or Negro populations. Their arms
are long in proportion to their legs, and their lower legs are
particularly short. Their heads are not small — Twiesselmann has
given cranial-capacity figures of 1,428 cc. for males and 1,268 cc.
for females in a series of eight skulls. Although this is not sta-
tistically impressive, eight is as many skulls as we have had to
work with in most fossil lines, and more extensive measurements
on the heads of living Pygmies confirm these figures.
Their faces are very short; their noses are short and wide, with
a nasal index, on the living, of 100 and more. In many cases their
eyes protrude, as if from exophthalmic goiter. Their hair is spiral,
as Negro hair is, but rarely peppercorn as among Bushmen. The
men have abundant beards, and many of them have hairy bodies,
with a particular abundance of chest hair. This is not fetal lanugo,
as some authors have claimed, but ordinary body hair. Many
Pygmy children have red hair, but this is caused by a nutri-
tional disease, kwashkior.
Although variable in skin color, the Pygmies are not black;
neither are they yellow like Bushmen. Their usual skin color is
dark reddish brown, which Gates calls mahogany. Their eyes are
dark brown, but the sclera is white, not flecked with melanin
patches as it is among many Negroes and Australian aborigines.
Little has been written about Pygmy teeth, except that they are
frequently carious and fall out early in life. Aside from this, we
know only that they are not small, like those of Bushmen.
I have studied several hundred photographs of Pygmies, but in
these I have seen only one man, Ilombe, chief of the Bambenga
in the northwestern Belgian Congo, who looks in the least Cauca-
soid, or Hamitic.2 Very few if any Pygmies resemble Bushmen or
Hottentots. This evidence suggests that the Pygmies began to
shrink before the ancestors of the Hamites and Capoids moved
southward; otherwise we would see more evidence among the
living Pygmies of the passage of these migrants.
1 agree with Gusinde and with Gates that the Pygmies are
descended from the old pre-Hamitic, pre-Capoid population of
the parklands and grasslands of Africa which was driven into the
forest by drought affecting both their water supply and their
2 Twiesselmann: op. cit.
Do the Pygmies Hold the Answer? 655
hunting. Once they were in the forest, one or more mutations for
dwarfing, which had already occurred among them outside the
forest, now acquired a survival value, and natural selection soon
spread this new trait through the forest populations. For our pur-
pose it does not matter whether all the Pygmies are descended
from one group of full-sized refugees or from several groups.
If we want to know what the full-sized ancestors of the Pygmies
looked like, all we need do is select a group of Pygmy children,
feed or inject them with the hormones the lack of which makes
them small, and see what they grow into. This is a perfectly feasi-
ble experiment and the Pygmies would probably co-operate.
In the meantime, we can reconstruct the image of a full-sized
man with a big body, a full-sized head, a broad face and broad
nose, eyes set wide apart, and probably heavy brow ridges, for the
Pygmies do not lack brow ridges. His skin was either mahogany-
colored or black ( the Pygmies may have become slightly depig-
mented in the forest), and his body was hairy. He may have had
convex, uneverted lips like the Pygmy instead of roll-out lips like
the Negroes. Such a man could well have been a descendant of
the Saldanha-Rhodesian-Cape Flats group and the ancestor of
both Pygmies and Negroes, the Pygmies, despite a reduction in
size, retaining the more archaic form.
Let us suppose that a population of these archaic, proto-Negro
and proto-Pygmy people comparable to those we have seen in
East and South Africa continued to live in West Africa well into
post-Pleistocene time, away from the path of Capsian and Ca-
poid migrations. Let us next suppose that such a population
mixed with Pygmies. By this mixture they would have acquired
the bulbous forehead, protruding eyes, and other infantile fea-
tures characteristic of living Negroes, features which distinguish
them from the Caucasoids whom their ancestors more closely re-
sembled.
To me this theory — that the modern Negroes resulted from a
backcross between an original proto-Negro stock and Pygmies,
which had evolved from the same ancestors by dwarfing — makes
sense. It explains the physical characteristics of the modern Ne-
groes, and it conforms with the evidence we have of their age, as
a race. In fact, their transformation need only have occurred a few
Africa
656
millennia before the historic expansion of Negroes over much of
Africa. And had modern Negroes existed very much earlier, some
would have wandered into East Africa and we might have seen
their remains.
This theory of the origin of Negroes does not exclude the pos-
sibility of mixture between proto-Negroes and Hamites or Ca-
poids. Such mixtures probably took place, and without doubt they
would explain some of the regional variations among Negroes.
But in this theory such mixtures are not a primary cause of the rise
of the Negroes. Because hybrids tend to return to one of their
parental stocks, no valid subspecies can arise through mixture.
Like the other four subspecies, the Congoids had an ancient, if
still little known, history.
Was Africa the Cradle of Mankind?
Darwin considered Africa to be the cradle of mankind. Later,
under the influence of Matthews, Osborn, and Andrews, the pen-
dulum of popular opinion swung to central Asia, where, we now
know, human beings were marginal and late. With the discovery
of Pithecanthropus, the cradle was thought to be southeast Asia,
and now Dart, Leakey, Arambourg, and others have again located
it in Africa.
It now seems likely that the Australopithecines evolved in Af-
rica, whence they spread to the east through the tropics of the
Old World. It is also possible, although it cannot be proved, that
the primary evolutionary step from Australopithecus to Homo
was taken, not on African soil, but in the Meganthropus-Pithecan-
thropus sequence. Java, and by extension all of southeast Asia, is a
serious rival.
Wherever Homo arose, and Africa is at present the likeliest con-
tinent, he soon dispersed, in a very primitive form, throughout the
warm regions of the Old World. Three of the five human sub-
species crossed the sapiens line elsewhere. If Africa was the
cradle of mankind, it was only an indifferent kindergarten. Europe
and Asia were our principal schools.
13
K
THE DEAD AND THE LIVING
T
_Lhe last four chapters are a unit. They constitute the
documentation for the racial history of man. The procession of
skull after skull with accompanying teeth and long bones may
seem a lengthy catalogue, but in reality it is not an overburden-
ing mass of evidence. A total of a little over three hundred bone-
bearing sites is not a large number: these sites encompass all our
knowledge about the ancestry of a species that now numbers over
two billions. All the pertinent information available in the litera-
ture on the subject and elsewhere had to be brought forth and
considered. Only by examining every scrap of data could I hope
to discover when and where each of the five lines of human de-
scent began, and where each led.
But before I could start on this documentation, I had to estab-
lish some degree of credibility for my thesis, which I state in
Chapter i. My thesis is, in essence, that at the beginning of our
record, over half a million years ago, man was a single species.
Homo erectus, perhaps already divided into five geographic races
or subspecies. Homo erectus then evolved into Homo sapiens not
once but five times, as each subspecies, living in its own territory,
passed a critical threshold from a more brutal to a more sapient
state.
This point of view is not wholly original — I know, for instance,
of two younger men who have thought it out independently of
myself and of each other 1 — nor is it generally accepted. As I was
working alone, with only Weidenreich’s interpretation of the Si-
1 Frank Livingstone of the University of Michigan and Loring Brace of the
University of California at Santa Barbara.
The Dead and the Living
658
nanthropus material to guide me, I decided, before writing this
book, to marshal many kinds of cognate evidence.
I studied genetic theory, zoogeography, and human physiology
(with special reference to adaptations to climate and culture);
the history of the primates, with its marvelous record of parallel-
ism, by which such similar creatures as the Old and the New
World monkeys could evolve from different prosimians; and the
record of our hominid predecessors, the Australopithecines. I also
made a survey of world archaeology covering the Pleistocene. In
addition, I had to explain the differences among fossil men be-
tween evolutionary characteristics and those that are racial.
These efforts filled eight chapters, numerically two thirds of the
book, but without them, Chapters 9 through 12 would not have
been solidly grounded.
Now that the task is over, I feel that the three Eurasiatic fines —
the Australoid, Mongoloid, and Caucasoid — have been traced
fully enough so that future discoveries will entail no major sur-
prises. The African material, however, is less well documented
and new conclusions may be reached as new evidence becomes
available.
As far as we know now, the Congoid fine started on the same
evolutionary level as the Eurasiatic ones in the Early Middle
Pleistocene and then stood still for a half million years, after
which Negroes and Pygmies appeared as if out of nowhere. The
Ternefine-Tangier fine has left us enough jaws and teeth to work
with, but no crania or cranial fragments except for one un-
measured parietal from Ternefine. Until a pre-Mouillian skull or
two are found in North Africa, a fair test cannot be given to my
facing page: This schematic map shows the distribution of the five subspecies of
Homo during most of the Pleistocene, from 500,000 to 10,000 years ago. This
distribution matches that on the diagram in Chapter 1. Of the five subspecies, the
Congoid was the most isolated; it was in contact with only one other, the Capoid,
then resident in North Africa. The second map shows what happened at the end of
the Pleistocene, when the Mongoloids and Caucasoids expanded and burst out of
their territories. The Mongoloids entered and inhabited America, and extended
their domain southward into Southeast Asia and Indonesia, while the Australoids
crossed Wallace’s Line and occupied Australia and New Guinea. The Caucasoids
thrust northward. More significantly, they drove the Capoids out of North Africa
and occupied the White Highlands of Kenya and Tanganyika. The Congoids were
reduced to a small part of their earlier domain, including the Congo forests and the
lands to the north, where they later evolved rapidly and spread, as Negroes, over
much of Africa.
66o
The Dead and the Living
hypothesis that the ancestors of the Bushmen and Hottentots
originated north of the Sahara and only reached South Africa
postglacially. The discovery of such a skull may also help ex-
plain why in the Middle Pleistocene North Africans resembled the
earliest Mongoloids, whereas the East Africans were closest to the
Caucasoids.
In addition to a Middle or Early Upper Pleistocene skull or two
from North Africa, we urgently need new evidence concerning
the details of the transition from the australopithecine to the hu-
man grade. The search for more early hominid fossils should
be accelerated in the few suitable areas of the Old World which
contain Lower Pleistocene deposits. Only when the key fossils
have been found will we know where and when the major lines
of human descent embarked on the separate paths that they have
followed to this day.
Toward the end of the Pleistocene, after all five geographical
races of man had become sapiens but before the two northern-
most, the Mongoloid and Caucasoid, had completed their south-
ward invasions and expansions, each race may have contained
nearly equal numbers of individuals. However, by the time agri-
culture and animal husbandry had been invented, by Caucasoids
and Mongoloids, these two had begun to outnumber the others.
With the wide spread of food production, the numerical dis-
proportion between the races increased; and today Mongoloids
and Caucasoids together constitute the vast majority of the
earth’s inhabitants.
The Australoids are on the decline, except among the aboriginal
tribes of India; and the Bushmen and Hottentots number only
tens of thousands. The Pygmies are few, but hold their own. The
African Negroes, on the other hand, have shown extraordinary
vitality. They have been particularly versatile in adopting new
cultures wherever they have been taken, as laborers, by Cauca-
soids and Mongoloids, and they have become the dominant racial
element in many of the tropical lowland regions of the New
World, as well as of Madagascar and parts of the Arabian coast.
Once a race has become established as the principal population
of a region, it has a tendency to stay there and to resist the genetic
influences swept in by later invasions. Less than a thousand years
66i
The Dead and the Living
ago the Arabs had a city near Amoy on the China coast, complete
with minarets and bazaars. Thousands of Arab men must have
impregnated Chinese women; yet today there is little if anything
about the Fukienese to show it. Kashmiri traders live, marry local
women, and die in the cities of Tibet, and Spaniards by the thou-
sands have settled in the Andean altiplano, but today Tibetans
and Andean Indians are as mongoloid as ever.
When two races come into contact and mixture occurs, one race
tends to dominate the other. The local advantage that the ge-
netically superior group (superior for its time and place) pos-
sesses may be primarily cultural or primarily physiological, or a
combination of both. For example, the dominance of the Euro-
peans over the native peoples of North America, Australia, and
New Zealand is primarily cultural; that of the Negroes in the
tropical lowlands of the New World and of the Indians in the
Andes is primarily physiological.
There is, however, a third kind of dominance, expressed by the
resistance of a population to the intrusion of large numbers of
outsiders into its social and genetic structures. Call it xenophobia,
prejudice, or whatever, people do not ordinarily welcome masses
of strangers in their midst, particularly if the strangers come with
women and children and settle down to stay. Social mechanisms
arise automatically to isolate the newcomers as much as possible
and to keep them genetically separate. This has happened his-
torically to Jews (who wanted to preserve their culture) nearly
everywhere, and to Negroes in the New World. It has happened
recently to Europeans in India and Indonesia, and in Africa it is
happening very dramatically to Europeans, even as I write.
The above is the behavioral aspect of race relations. The ge-
netic aspect operates in a comparable way. Genes that form part
of a cell nucleus possess an internal equilibrium as a group, just
as do the members of social institutions. Genes in a population
are in equilibrium if the population is living a healthy life as a
corporate entity. Racial intermixture can upset the genetic as well
as the social equilibrium of a group, and so, newly introduced
genes tend to disappear or be reduced to a minimum percentage
unless they possess a selective advantage over their local counter-
parts.
662
The Dead and the Living
I am making these statements not for any political or social
purpose but merely to show that, were it not for the mechanisms
cited above, men would not be black, white, yellow, or brown.
We would all be light khaki, for there has been enough gene flow
over the clinal regions of the world during the last half million
years to have homogenized us all had that been the evolutionary
scheme of things, and had it not been advantageous to each of the
geographical races for it to retain, for the most part, the adaptive
elements in its genetic status quo.
This status quo entails not only the variations in bones and
teeth that are evident in fossil man, and those of the surface
features of living men, like skin, hair, lips, and ears, by which
we can distinguish races almost at a glance, but also subtler dif-
ferences seen only on the dissecting table or through the eyepieces
of microscopes. Races differ in the extent and manner in which the
fine subcutaneous muscles of the lips and cheeks have become
differentiated from the parent mammalian muscle body; in the
chemical composition of hair and of bodily secretions, including
milk; in the ways in which different muscles are attached to bones;
in the sizes and probable secretion rates of different endocrines;
in certain details of the nervous system, as, for example, how far
down in the lumbar vertebrae the neural canal extends; and in
the capacity of individuals to tolerate crowding and stress.
These and other details of racial difference I hope to describe
and document in a later volume.
In studying racial differences in living men, physical anthro-
pologists are now relying less and less on anthropometry and more
and more on research in blood groups, hemoglobins, and other
biochemical features. This is all to the good because the inherit-
ance of these newly discovered characteristics can be accurately
determined. In them, racial differences have been found, differ-
ences just as great as the better known and much more conspicu-
ous anatomical variations. Being invisible to the naked eye, they
are much less controversial than the latter in an increasingly race-
conscious world. To me, at least, it is encouraging to know that
biochemistry divides us into the same subspecies that we have
long recognized on the basis of other criteria.
To readers who find these simple biological facts disconcerting,
The Dead and the Living 663
let me repeat something I said in Chapter 1. Until the present
century, and in some countries until the present day, all five sub-
species of man whose racial histories I have traced include popu-
lations of food gatherers and hunters living in the same regions
that their ancestors occupied at least as early as early postglacial
times. Some of the most backward in a cultural sense belong to the
Mongoloid and Caucasoid subspecies, other populations of which
have achieved the highest levels of civilization yet known in the
world. But these backward populations do not live in their ances-
tral homelands; they hunt in distant regions that their ancestors
invaded.
Caucasoids and Mongoloids who live in their homelands and
in recently colonized regions, such as North America, did not rise
to their present population levels and positions of cultural domi-
nance by accident. They achieved all this because their ancestors
occupied the most favorable of the earth’s zoological regions,
in which other kinds of animals also attained dominance during
the Pleistocene. These regions had challenging climates and
ample breeding grounds and were centrally located within con-
tinental land masses. There general adaptation was more impor-
tant than special adaptation. Any other subspecies that had
evolved in these regions would probably have been just as suc-
cessful. Now the success of these groups is being challenged in
many parts of the world as other groups who evolved later learn
to use their inventions, especially modern means of communica-
tion. And evolution is still taking place, particularly natural se-
lection resulting from crowding and stress, as described in Chap-
ter 3.
In any case, neither the future of man nor the detailed descrip-
tion of the bodies, biochemical peculiarities, or behavior patterns
of the living races of man is the subject of this book. I have, I hope,
shown as accurately as the evidence warrants whence each of
them came, and what steps guided it to its present position.
Further details must await the publication, in due course and if all
goes as planned, of my next book, tentatively entitled The Living
Races of the World.
STATISTICAL APPENDIX
BIBLIOGRAPHY
GLOSSARY
INDEX
TABLE 36
ARCS AND CHORDS OF THE FRONTAL,
PARIETAL, AND OCCIPITAL BONES
IN THE SAGITTAL PLANE
Frontal Parietal Occipital
Skull
Arc
Chord
Index
Arc
Chord
Index
Arc
Chord
Index
Pithecanthropus 1
(100)
98
(98.0)
91
87.5
96.0
(103)
78
(75.7)
Pithecanthropus 2
90?
88
97.7?
94
91
95.8
101?
75
74.2
Sinanthropus 2
123
113
91.8
112
104
93.1
Sinanthropus 3
115
102
90.5
100
94
94.0
106?
80?
74.2?
Sinanthropus 10
129
115
89.2
113
106
94.0
Sinanthropus 11
122
106
86.9
92
86
93.5
118
86
72.9
Sinanthropus 12
124
113
91.2
102.5
91
95.7
118
86
72.9
Solo 1
139
120.5
86.5
106
101
95.5
111
81.5
74.1
Solo 5
136
120
88.3
117
111
95.2
128?
94?
73.3
Solo 6
122
112
91.7
107
102
95.6
109
82
75.2
Solo 9
103?
99?
96.3
115?
88
76.4
Solo 10
135
120
89.0
105
102
97.3
114
78
68.4
Solo 11
122
112
91.8
102
97
94.2
122
90
73.4
Mapa
134
115.6
86.3
114
107
87.2
(109.0)
(87.2)
(79.9)
Liu-Kiang
136.5
117.2
85.9
117.2
119.2
91.5
105.5
91.5
86.7
Tze-Yang
103?
92?
89.0
116
104
90
98?
84
86
Wadjak 1
136
119
87.5
130
113
86.9
Rhodesian
137.5
121
88.0
117
112
96.0
118
b9
74.1
Steinheim
118
110
84.7
103
96
93.2
(117)
(90)
(76.9)
Swanscombe
118
109
92.3
118
95
80.5
Ehringsdorf
135
115
85.4
128
119
93.2
117
87
74.3
La Chapelle
121
107
88.5
121
112
92.5
115
91
79.1
La Ferrassie 1
125?
114?
91.2
114?
106?
93.0?
124
99
79.8
Neanderthal
133
116
87.3
110
104
94.7
Spy 1 M
100.0?
102.8?
93.4?
126.0?
114.9?
91.2
Le Moustier
120.2?
108.2
90.0?
121.8
109.2
96.3?
(adolescent)
Gibraltar 1 F
124.0?
107
86.3?
106.0
81.1
76.5
La Quina 1 F
116.3
106.4
91.5
106.9
102.9?
96.3?
Spy 2 F
115.0
109.0
94.8
Combe Capelle
138
123
89.1?
132?
123?
93.2?
128?
100?
78.1?
Cro-Magnon
144
119
82.6
138
123
89.1
128
109
85.1
Grimaldi
135
115?
85.2?
145?
131
90.3
130?
97?
74.6?
TABLE 37 ( continued )
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TABLE 37 ( continued )
F. MOUILLI ANS AND CAPSIANS; NORTH AND EAST AFRICA
Taforalt Gamble’s Cave Naiva- Olduvai Elmenteita
(14) (13) 4 5 sha R. R. Bed 5 A B C D
Sex (± Age) MF MM M M MMMYF
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TABLE 38
DIMENSIONS AND INDICES OF MANDIBLES
Pithe- Unner
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(Symphysial) 70° 62.5° 63.5° 63° 73° 74° 65°
Mandibular (Gonial) Angle 118° 110° 109°
TABLE 38 ( continued )
Tabun Skhul Ternefine Sidi Abd Temara Rabat Fish Asselar
TABLE 39
I2 is larger than I1 because teeth are from different individuals. L = length, B ==» breadth, R =* robusticity
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6.2- 9.0
5.0- 9.0
5.0- 8.5
5.8- 9.3
7.0-11.0
5.5- 9.5
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125
Broken
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Mean
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TABLE 39 ( continued )
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9.7 10.0-11.0
11.3 11 10.0 4.0-15.0
11.5 10.5 8.0 4.0-13.0
13.0 115.5 80
BIBLIOGRAPHY
Abbie, A. A.: “The Quest for Man’s Birthplace.” AuS, Vol. 1, No. 4 (1961), pp.
201-7.
Adam, W.: “The Keilor Fossil Skull, Palate, and Upper Dental Arch.” MNMM,
No. 13 (1943), pp. 71-8.
Adams, T., and Covino, B. G.: “Racial Variations to a Standardized, Cold Stress.”
JAP, Vol. 12, No. 1 (1957), pp. 9-12.
Agache, R., and Bordier, F.: “Decouverte de Silex Apparemment Tailles a un
Equide Archaeique de Type Villafranchien dans la Haute Terrasse Superieure
de la Somme.” CRAS, Vol. 248, No. 3 (1959), pp. 439-40.
Alcobe, S.: “Die Neanderthaler Spaniens.” NC, 1958, pp. 9-62.
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NOTICES
Discovery of Fossil Skeletons of Small People in a Cave on the Island of Flores,
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GLOSSARY
acetabulum The hip socket.
achondroplastic A form of dwarfing in which the extremities are
shortened and thickened.
acrocentric See telocentric.
Adapidae Fossil ancestors of the lemurs and lorises.
adduction A drawing together, as the adduction of the great toe toward
the other toes in human evolution.
adductor magnus A large muscle extending from the pelvis to the linea
aspera of the femur. It is important in locomotion.
alleles Alternate genes situated on a single locus of a chromosome.
allometry (allometric) The principle according to which the propor-
tions of related animals change as their sizes change.
allop atric Inhabiting different regions.
alveolar Pertaining to the tooth-bearing part of the upper jaw.
anagenesis The evolution of one species out of another by succession:
phyletic evolution.
Anaptomorphs ( Anaptomo rphidae ) Fossil tarsiers.
anastomosis A connecting link between two arteries which ensures
blood flow to the vascular territories of both if one is cut.
anorthoclase A feldspar.
annulus A bony ring around the ear hole in certain mammals.
anteroposterior Fore and aft.
anticlinal (vertebra) In jumping primates, the vertebra that marks
the midpoint between the front and rear muscles of the back on a pono-
grade animal.
apocrine (gland) A kind of sweat gland.
aponeuroses Tendinous sheaths.
Archaeozoic The oldest of the five geological eras.
argon-potassium method, or Argon-40 method. A method of geological dat-
ing based on measuring the amount of argon trapped in a potassium
atom. See page 313.
asterion The point on the surface of the skull where the lambdoid,
parietomastoid, and occipitomastoid sutures meet.
astragulus The ankle bone.
ateliotic A kind of dwarfing in which the bodily proportions remain
normal with due allowance for allometry.
Aterian A North African flint industry characterized by bifacial pressure
flaking and tanged.
712 Glossary
atlas (vertebra) The second cervical.
Aurignacian A European Upper Paleolithic flint industry.
Australoid One of the five subspecies of living man, including the
native peoples of Australia, New Guinea, and Melanesia; the Negroid
dwarfs of Indonesia and South Asia; and certain aboriginal tribes of
India.
Australopithecines All Lower Pleistocene hominids that are not Homo.
autosomal cells Body cells, as opposed to sperm and egg cells.
AXILLA Armpit.
basal conglomerates The conglomerate (puddingstone) which often
occurs at the base of beds deposited on a surface that was exposed and
eroded before the deposition of the new series.
basion The point at the forward lip of the foramen magnum on the
sagittal line.
biasterionic The breadth of the skull on the chord between the left and
right asterion (q.v.).
bicondylar (diameter) The chord between the two condyles of the
mandible, usually taken between the outer borders of the condyles.
bicristal diameter The distance between the crests of the ilia of the
two pelvic bones.
bigonial diameter The length of the chord between the two gonial
angles of the mandible.
bizygomatic diameter The maximum face breadth measured from
one zygomatic arch to the other.
bilophodontism A feature of the molar teeth of Old World monkeys.
Each molar has two pairs of cusps, one forward and the other to the
rear, each pair being joined by a ridge to form a loph.
biogeography The geography of living things, including both plants and
animals.
bolas stones Stones tied together with thongs so that when thrown at
an animal or bird they will spread in flight and entangle the victim.
boreal Pertaining to the northerly regions; e.g., boreal forest.
brachial Pertaining to the upper arm; e.g., the brachial plexus of
nerves.
brachiate To move through the forest by swinging with the arms from
branch to branch, as an ape does.
brachycephalic Round- or short-headed.
brachycrany Of a skull: short or round.
branching (evolution by) The evolution of one or more sister species
at the same time through environmental adaptation: kladogenesis.
breccia A cave deposit containing angular stone and bone objects that
have been naturally cemented.
bregma The point at or near the top of the skull where the frontal and
parietal sutures meet.
buccal On the cheek, or outer, side of the teeth.
bulla (tympani) A hollow, thin-walled, bony prominence of rounded
form situated just below the opening of the ear and forming part of the
tympanic bone.
Glossary
7i3
burin A graver or narrow chisel, particularly of flint.
calcaneus The heel bone.
calotte A small, caplike fragment of a brain case.
calva A fragmentary brain case, usually lacking the base.
calvarium The brain case, without the face.
capitatum (os capitatum) A wrist bone.
Capoid One of the five subspecies of man, including the Bushmen and
ffottentots.
Carabelli’s cusp An accessory cusp On the lingual side of an upper
molar tooth.
carpal (bones) Wrist bones.
carotid An artery supplying blood to the brain.
Catarrhines Old World monkeys, apes, and hominids.
Caucasoid One of the five subspecies of living man, including most
Europeans, North Africans, Near Easterners, inhabitants of India and
Pakistan, and overseas settlers from these regions.
Cenozoic The present geological era, including the Tertiary and Quater-
nary.
centromere The point where the two coiled strands of a chromosome
are attached; in cell division it acts as the focus of separation.
cerebellum A three-lobed and finely fissured section of the brain situ-
ated below the posterior portion of the cerebral hemispheres. Among its
functions are the regulation of posture and the maintenance of bodily
equilibrium.
cingulum A collarlike rim of enamel about the base of the crown of any
tooth except the incisors.
Clactonian A European Lower Paleolithic flake industry.
class The third of seven levels in the Linnean taxonomy; e.g., Mam-
malia.
clavicle Collarbone.
CLINE, clinal A gradual progression in the dimensions, form, or color
of an anatomical feature from one geographic region to another.
condyles Raised articular surfaces on which bones move, as the oc-
cipital condyles at the base of the skull, and the condyles of the lower
jaw.
Congoid One of the five subspecies of living men, including the African
Negroes and Pygmies.
coracoid process The forward peak of the ascending ramus of the man-
dible to which the temporal muscles are attached.
cortex The outer layer of the lobes of the forebrain; the “gray matter.”
cretaceous The third and last epoch of the Mezozoic Era.
Cro-Magnon A site in the Dordogne region of France in which a skele-
ton so named was found. That skeleton. A supposed race.
Cromerian A Lower Pleistocene interglacial period between Giinz and
Mindel.
cuboid A tarsal bone.
cuneiforms Three tarsal bones.
7M
Glossary
cynodont, cynodontism Being “dog-toothed.” Having a normal-sized
pulp cavity: the opposite of taurodontism (q.v.).
cytogenetics The study of heredity in terms of the anatomv and physi-
ology of cells.
deciduous teeth The milk teeth.
deltoid A muscle of the shoulder and upper ann.
diastema A gap between two teeth.
digastric fossae Grooves on the inner and under side of the lower jaw
to which the digastric muscles are attached.
dimorphism (sexual) Marked differentiation in size and form between
the sexes.
diploe The cancellous bony tissue between the inner and outer tables of
the skull.
diploid Of cell nuclei: having pairs of chromosomes; e.g., 46 in man.
The opposite of haploid ( q.v. ) .
distal Away from the center; e.g., the hand is distal to the arm.
Djetis A late Lower Pleistocene fossil-bearing deposit found in Java.
dolerite A dark igneous rock.
Donau A local Central European Lower Pleistocene glaciation.
dorsal On the back side.
Dryopithecines A group of fossil hominoids probably ancestral to living
apes and men.
Elster The first European continental icecap, corresponding to the
Mindel of the Alpine series.
ecology The study of the mutual relations between organisms and
their environment.
Eocene The second of five divisions of the Tertiary epoch.
eoliths Early stone “implements” which may have been of natural
origin.
epoch (geological) A division of a period (q.v.).
era (geological) Any of the five primary divisions of geologic time.
estivate To sleep in the hot season— the opposite of hibernate.
Ethiopian region Africa south of the Sahara, and Southern Arabia.
euryphagous, euryphagy Wide-feeding, eating many kinds of food.
exfoliate To peel off through weathering (of granite).
exostosis A bony excrescence.
family The fifth of seven levels in the Linnean taxonomv; e.g.,
Hominidae, which has two subfamilies, Australopithecinae and
Homininae.
feldspar Any of a related group of crystalline minerals.
femur Thigh bone.
fibula The thinner and outer of the two shin bones.
flake (implement) A stone implement made from a flake that has been
struck off a core.
foramen A hole in a bone.
foramen magnum The opening at the base of the skull through which
the spinal cord passes.
fossa A depression or concavity in a bone.
Glossary
715
fovea An indented area in the center of the retina.
frenum A connecting fold of membrane in the mouth. One connects
the lips and gums at the median line, and another binds the tongue
to the floor of the oral cavity.
Gamblian The last (either third or fourth) pluvial period of the
Pleistocene in East Africa.
general adaptation Genetic adaptation to interspecific and intra-
specific competition; e.g., increased intelligence.
genetic drift (the Sewall Wright effect) A type of evolution postulated
by Sewall Wright in which the genetic composition of a population
changes by chance because the population is too small to constitute a
valid statistical sample.
genioglossal A pair of muscles that participate in controlling the
movements of the tongue.
genus The sixth of seven levels in the Linnean taxonomy; e.g., Homo.
geographical race A subspecies; a major division of a species, pre-
sumably of some antiquity.
gerontomorphic, gerontomorphy Having the characteristics of old
age, as contrasted with pedomorphic (q.v.).
gibbehellic acid A substance involved in the growth of plants.
glabella The central and most prominent point on the brow ridges.
glabellare An ill-defined point on the sagittal arc of the frontal bone
just above glabella, marking the junction of the glabellan prominence
and the curve of the frontal bone.
glacial An interval of cold climate with mountain glaciers, continental
icecaps, or both.
glenoid cavity The socket in which the mandibular condyle rests.
glottochronology The science of determining how long ago two re-
lated languages separated.
gluteus maximus A massive muscle of the pelvis and thigh which
forms the buttock. It is important in walking and particularly in raising
the body upward.
gluteus medius A powerful muscle of the pelvis and thigh. In walk-
ing it throws the body toward the line of gravity when the opposite leg
is off the ground.
gluteus minimus A muscle of the pelvic and thigh underlying the
gluteus medius and performing more or less the same function.
gonial angle The lower rear corner of the mandible.
Gottweig The interstadial between Wiirm I and Wiirm II.
grade In this book, an evolutionary level or status through which one
or more phyletic lines of animals (or plants) may pass.
Gunz The first of four Alpine glaciations in Europe; believed to have
been divided into two peaks, Giinz I and Gunz II.
haploid The condition of a sperm cell or unfertilized egg which has
only one pair of chromosomes; e.g., in man, 23, which is half the num-
ber present in fertilized eggs and in somatic cells.
haptoglobins “Proteins [in the blood serum which are] concerned with
the binding [together] of hemoglobin from aged and broken down red
blood cells” (C. Stem: Principles of Human Genetics, p. 53).
7i 6 Glossary
hemoglobin “Complex molecules [in the red blood cells] that are com-
posed of the colored, iron-containing heme and a colorless protein,
globin’’ (Stern: Principles, p. 53).
holotype The first specimen of a species to be found, named, and
described.
Hominids Australopithecines and men.
Hominoids Apes, Australopithecines, and men.
Homo erectus The extinct species of man from which the living races
of Homo sapiens evolved.
Homo sapiens The living species of man and some of our fossil ancestors.
humerus The upper arm bone.
Hylobatidae, Hylobatids Gibbons and siamangs.
hyoid bone A U-shaped bone at the base of the tongue; to it are at-
tached muscles used in swallowing and speaking.
hypophysis The pituitary gland.
hypothalamus A small subchamber at the upper end of the brain
stem which has many functions concerned with emotions and automatic
controls of physiological activities.
ilium The uppermost and largest of the three pelvic bones which fuse
to form the os coxae.
inca (bones) Supernumerary bones at lambda.
infraorbital foramen A foramen in the zygomatic bone under each
orbit.
inion A projection in the sagittal line of the occipital bone, usually at or
below the hindmost point of that bone. It serves as an anchor for some
of the neck muscles.
interglacial An interval of warm climate between two glaciations.
interpluvial In tropical regions, a geological time interval separating
two pluvials.
interstadial A cool interval between two maxima of a single glaciation.
ischium ( ischial ) The lowermost and hindmost of the three pelvic bones
which fuse to constitute the os coxae.
ischial callosity A patch of tough, bare skin covering the tuberosity
of the ischium in Old World monkeys and some apes.
ischial tuberosity A downward extension of the ischium, covered in
some primates by an ischial callosity (q.v.) .
Jurassic The second of the three epochs of the Mesozoic Era.
karyotype The chromosomes of an animal arranged by pairs in order of
length.
kingdom The first of seven levels in the Linnean taxonomy; e.g., the
animal and vegetable kingdoms.
kitchen-midden An open-air archaeological site, usually Mesolithic, and
usually containing bivalve shells.
kladogenesis Evolution by branching (q.v.).
labiolingual diameter The transverse diameter of a tooth, from
cheekside to tongueside.
lagomorphs Hares and rabbits.
Glossary 717
lambda The point where the two parietal bones and the occipital bone
meet.
lanugo Fetal hair.
laterite A porous reddish clay formed by the decomposition of certain
rocks in tropical regions.
Levalloisian A technique of striking flakes ready for use from a pre-
pared core. A flint-tool industry based on this technique.
line In this book, an evolutionary sequence of species passing through
two or more grades.
linea aspera A raised ridge on the back of the femur.
local race A minor taxonomic division of a species of lesser magnitude
than a subspecies (geographical race) .
loph In the molar teeth of Old World monkeys, a pair of cusps joined
by a ridge. There are generally two pairs to a tooth, one fore and one
aft.
lumbar The region of the back between the rib cage and the pelvis;
e.g., lumbar vertebrae.
malar The os zygomaticum, or cheek bone.
mandible Lower jaw.
mano (American Indian) The upper grindstone, held in the hand and
rubbed against the metate, or nether grindstone.
manubrium The lower segment of the breast bone.
marl A crumbly deposit of clay mixed with calcium carbonate.
masseter One of a pair of muscles which take part in the raising and
grinding motions of the lower jaw.
mastoid process A hollow protuberance on the temporal bone behind
the ear hole.
maxilla The upper jaw bone.
auditory meatus The ear hole.
medulla (oblongata) The lowest or posterior part of the brain,
which merges into the spinal cord.
meiosis A process by which a sperm or egg cell divides twice in succes-
sion, the second division reducing the nucleus from the diploid (q.v.)
to the haploid ( q.v. ) state.
melanin Pigment granules.
mental foramen A hole in the outer surface of the lower jaw.
meningeal arteries Arteries that feed blood to the meningeal cov-
ering of the brain.
mesial Toward the center.
mesiodistal diameter Of the crowns of teeth: the length as measured
along the curve of the dental arc.
mesiolabial Of the crowns of teeth: forward and outward.
Mesozoic The fourth of the five geological eras.
metacarp als The five bones of the hand lying between the wrist bones
( carpals ) and the finger bones ( phalanges ) .
metacentric Of a chromosome: having the centromere more or less in
the middle.
?i8
Glossary
metatarsals The five bones of the foot lying between the tarsals and
the toe bones (phalanges) .
metate An American Indian millstone, the larger or nether stone.
metopic suture A suture between the two frontal bones which usually
closes in infancy.
microlith A small flint implement struck from a small blade core.
midden An ancient refuse heap, usually composed largely of mollusc
shells.
mid-phalangeal Pertaining to the middle section of a finger or toe.
Mindel The second of four Alpine glaciations of the Pleistocene.
Miocene The fourth of five epochs of the Tertiary period.
Mongoloid One of the five subspecies of living man, including prin-
cipally the peoples of East and Southeast Asia and most of Indonesia,
and the Polynesians, Micronesians, and American Indians.
monotreme An egg-laying mammal of Australia.
monotypic Of a species: having no geographical races.
morphology The study of the form and structure of animals and plants.
mosaic A kind of geographical distribution in which many races live
close together.
Mousterian A Middle Paleolithic flake industry of Europe and Western
Asia.
muller A stone used for pounding or grinding.
multituberculates An extinct order of mammals that flourished dur-
ing the early Cenozoic.
nasion The point where the two nasal bones and the frontal bone meet.
navicular A wrist bone.
neanthropic Referring to morphologically modern types of man.
Nearctic region North America and parts of Central America (see
Map 2) .
neoteny The persistence into adult life of fetal or infantile character-
istics.
Neotropical region South America and parts of Central America (see
Map 2) .
neurone A nerve cell, including its processes.
nonton al (language) A language in which tones have no semantic
meaning, other than for emphasis or questioning.
Notopuro Upper Pleistocene fossil-bearing beds in Java.
nuchal Pertaining to the neck.
obelion, obelionic The point on the interparietal (sagittal) suture of
the skull between the two parietal foramina.
occiput, occipital bone The hindmost bone of the skull.
occipital crest A crest of bone running horizontally across the occiput
in some apes and some hominids.
occlusal Pertaining to the occlusion of the teeth when the jaws are
closed.
odontology The study of teeth.
Glossary
719
olecranon process The projection at the proximal end of the ulna: the
“funny bone.”
Oligocene The third of five divisions of the Tertiary epoch.
ontogeny The growth and development of an individual organism.
opisthion The rearmost point on the sagittal line of the skull when in
the eye-ear plane.
order The fourth of seven levels in the Linnean taxonomy; e.g., Pri-
mates.
Oriental region The Old World tropics from India to Wallace’s Line
and South China (see Map 2).
orthognathous Having jaws that do not protrude: the opposite of
prognathous.
os capitatum A wrist bone.
os coxae The pelvic bone, consisting of three fused bones, the ilium,
ischium and pubis,
os lunatum A wrist bone.
pachyostosis Thickening of a bone.
Palearctic region Europe, North Africa, and all of Asia except that
forming part of the Oriental region (q.v.). See Map 2.
Paleocene The earliest of the five divisions of the Tertiary epoch.
Paleolithic The stone-age industries of the Pleistocene and their time
span.
Paleozoic The second of the four geological eras.
parietal One of a pair of bones comprising the major part of the skull
vault, bordered in front by the frontal bone and behind by the occipital
bone.
patella The kneecap.
pedomorphic, pedomorphy Infantile or childlike in the adult form.
pelage The over-all hair covering or fur of an animal.
period (geological) A division of an era (q.v.).
peroneal Pertaining to the fibula, as the peroneal artery.
phalanges Finger and toe bones.
phenotype What you are: the product of heredity and environment.
phratry A division of a tribe or other breeding isolate.
phyletic evolution Evolution by succession by which one species
evolves out of another.
phylogeny The evolution of a line (q.v.).
phylum The second of seven levels in the Linnean taxonomy; e.g., the
Chordata- Vertebrata is a sub-phylum.
phytogeography The geography of plants; the counterpart to zoo-
geography (q.v.).
pilaster (femur) A bracing ridge on the back of the shaft of the thigh
bone.
Pin j or A Lower Pleistocene fossil-bearing geological formation in the
Siwalik Hills of India, laid down during the first of four Himalayan
glaciations.
720 Glossary
Platyrrhines New World monkeys.
platysma A broad, thin sheet of muscle covering much of the face in
primitive mammals; from parts of it are derived the muscles of facial
expression in man.
Pleistocene The earlier and longer of the two epochs of the Quaternary
period; it began about one million years ago and is believed to have
ended ten thousand years ago.
plexus A network of blood vessels or nerves.
Pliocene The fifth and final epoch of the Tertiary period.
pluvial Rainy. In Africa pluvial periods are believed to have corre-
sponded in a general way to periods of glaciation in northerly regions.
polytypic Of a species: having two or more genetically distinct geo-
graphical races or subspecies.
polyandrous Pertaining to a system of mating in which a woman may
have two or more husbands.
polygynous Pertaining to a system of mating in which a man may have
two or more wives.
polymorphic Genetically variable.
pongid Pertaining to apes.
Pontian A geological time span between about 16 and 10 million years
ago, variously attributed to the Upper Miocene and the Lower Pliocene.
porion A point on the upper border of the ear hole.
premolar cone A protuberance in the center of an upper premolar
found sometimes in the teeth of Mongoloids.
prognathism Protrusion of the jaws.
pronograde Walking on all fours.
Prosimian Any primitive primate included among the tree shrews,
lemurs, lorises, and tarsiers.
Proterozoic The second of the five geological eras.
protoanthropic The first of three grades of human evolution accord-
ing to the scheme of S. Sergi.
pterygoids Two pairs of muscles, the internal and the external, con-
cerned with the motions of the lower jaw.
purine A crystalline compound (C2H4N4), the parent of other com-
pounds of the uric acid group.
Quaternary The present geological period, including the Pleistocene
and Recent epochs.
race A general term referring to genetically distinct divisions of a
species.
radius The shorter and more mobile of the two bones of the forearm;
the one on the outer side when the palm of the hand is facing forward
or upward.
rami (ascending) The paired branches of the rear portions of the lower
jaw which rise upwards to articulate with the cranium.
rete mirabile A network of blood vessels concerned with heat trans-
fer between veins and arteries.
Rhesus An obsolete name for the primate genus Macaca; a system of
blood groups named after the Rhesus monkey, Macaca mulata.
Glossary y21
Rhinial The name of the third index of facial flatness.
Riss The third of the four Alpine glaciations of the Pleistocene epoch.
Saale The second Continental icecap in Northern Europe, correspond-
ing to the Riss Alpine glaciation.
sacrum A bone of composite origin connecting the two pelvic bones,
the fifth lumbar vertebra, and the coccyx.
sagittal CREST A crest running along the sagittal (interparietal) suture
in some apes and Australopithecines whose two sets of temporal mus-
cles are or were so large that they met at the top of the skull and
needed additional area for attachment.
Sahul Shelf A region of shallow water off the northwest coast of
Australia which reaches New Guinea; it was dry land during parts of
the Pleistocene. & 1
Sanmenian Geological deposits in Northern China: the lower San-
menian is Lower Pleistocene and the Upper Sanmenian is Middle
Pleistocene.
SAVANNAH
SCAPHOID
SCAPULA
Tropical or semitropical grassland dotted with trees.
Keel-shaped: a wrist bone.
Shoulder blade.
sclera The fibrous outer capsule of the eye, including the transparent
cornea. r
sectorial Of canine and incisor teeth: shearing.
SELLA turcica The hypophysial fossa, a depression in the base of the
skull in which the pituitary or hypophysial gland is seated.
serval CAT ( Felis Serval) A long-legged African wildcat.
sesamoids Generally rounded or platelike bones found in tendons over-
- 11 8 joints. The patella or kneecap is a large sesamoid bone.
shaman A medico-religious specialist among primitive peoples.
shovel incisors Incisor teeth that are concave on the inside.
sickling A heritable deformation of the red corpuscles which inhibits
oxygen transfer but is believed to produce immunity to some forms of
malaria.
Simotic
The second of the four indices of facial flatness.
Sinanthropus A group of fossil men found at Choukoutien in North
China.
special adaptation Genetic adaptation to some special environmental
factor; e.g., cold, heat, drought, or special foods.
species The seventh of the seven levels in the Linnean taxonomy, and
the basic unit of the Linnean system; e.g., Homo sapiens.
sphenoid A bone in the lower and forward part of the skull. It consists
of a body and two pairs of wings and articulates with every other bone
of the brain case as well as with the palatine and zygomatic bones and
the vomer.
squamous Pertaining to the upper or scalelike part of the temporal bone
which articulates with the parietal at the squamous suture.
stadial One of the maxima of a glaciation which had two or more
peaks.
\)
722 Glossary
steatopygia The condition of having large deposits of fat on the but-
tocks.
stegodon A genus of extinct Asiatic elephants.
stenophagous, stenophagy Having a specialized diet.
sternum The breast bone.
stratigraphy, stratigraphic The study of superimposed layers
(strata) in archaeology.
subfamily See family.
subphylum See phylum.
subspecies A major division of a species, constituting a geographical
race.
subtense In an isosceles triangle, a line that divides it into two right-
angle triangles of equal area.
subterminal Of a chromosome: having its centromere located between
its middle and one end.
succession (evolution by) See anagenesis, phyletic evolution.
Sunda Sea, Shelf A body of shallow water lying between parts of
Indonesia and Southeast Asia. During parts of the Pleistocene this area
was above water.
supersternale A point at the top of the sternum or breast bone used
in the anthropometry of the living.
supinator ridge or crest A ridge on the inside of the ulna which gives
attachment to the supinator muscle, which rotates the forearm.
sustentaculum tali A projection on the inner side of the calcaneum
or heel bone for articulation with a facet of the head of the talus, or
ankle bone.
suture The line of union between any two bones of the skull.
sylvian fissure Also called the lateral sulcus. The most conspicuous
fissure of the brain, situated between the temporal lobe and the fronto-
parietal region.
symbiosis Of two or more species: living together to the mutual ad-
vantage of both or all species; e.g., a bird that sits on a rhinoceros and
eats his ticks.
sympatric Of species: occupying the same territorv.
symphision An anthropometric landmark at the upper and outer border
of the pubic symphysis.
pubic symphisis The junction of the two pubic bones that are separated
by the interpubic disc.
taiga A Russian term for boreal forest.
talus The ankle bone, which articulates with the tibia, fibula, calca-
neum, and navicular. In walking each talus bears in turn the entire
weight of the body.
tarsal bones or tarsus The talus, calcaneum, navicular, cuboid, and
the three cuneiforms.
Tatrot An Early Lower Pleistocene level in the Siwalik Hills of North-
ern India, and its fauna.
taurondontism Being “bull-toothed.” A condition of the teeth, particu-
Glossary
723
larly molars, in which the fusion of the roots neotenously takes place
low down and results in a large pulp cavity. The opposite condition is
cynodontism (q.v.).
taxonomy The science of classifying animals and plants; systematics.
tektite A small, glassy nodule from outer space.
telocentric Of a chromosome: having the centromere at one end;
acrocentric.
temporal BONE A paired bone forming part of the base and lateral wall
of the skull. It is formed from four separate bones in the fetus, repre-
senting the petrous portion and mastoids, the styloid process, the
squamous part, and the tympanic part.
TERES minor A muscle attached to the axillary border of the scapula
and the upper part of the humerus, which participates in rotating the
humerus laterally.
Tertiary The first of the two periods of the Cenozoic Era.
Tiglian A cool interglacial interval of the Lower Pleistocene in Western
Europe preceding Giinz I.
tonal (language) A language, like Chinese, in which changes of
musical tone have semantic meaning.
torus A bony ridge, particularly the supraorbital torus, which is a
continuous brow ridge.
trapezium A wrist bone at the base of the first (thumb) metacarpal.
trapezoid A wrist bone at the base of the first (index) finger meta-
carpal.
travertine Calcium carbonate deposited by water of springs and
streams holding lime in solution.
Triassic The first period of the Mesozoic Era.
Trinil Fossil-bearing Middle Pleistocene beds in Java overlying the
Djetis Beds and underlying the Notopuro Beds.
trochanter At the upper and outer side of the femur, just distal of the
neck, is a large prominence, the greater trochanter, to which are at-
tached the gluteus minimus and gluteus medius muscles. The lesser
trochanter, lower and on the inner side, is the attachment for the psoas
major and iliacus muscles. A few femora have a third trochanter.
trochlea A front-to-back rounded groove at the distal end of the
humerus which articulates with the ulna.
tympanic plate A curved plate on the temporal bone which lies in
front of the mandibular fossa and with which the condyle of the
mandible articulates in rotary chewing.
ulna The longer and less mobile of the two bones of the forearm- the
one on the inner side when the palm of the hand is facing forward or
upward.
ungulates Hoofed mammals.
venae comites Veins which run parallel to each other on either side
of certain arteries and which may be joined by anastomoses (q.v.)
just as arteries may. ^
Villafranchian The earliest part of the Lower Pleistocene.
724
Glossary
volar Referring to the palm of the hand or sole of the foot.
vomer A median bone which forms the posteroinferior portion of the
nasal septum. It lies below and behind the septal cartilage.
Waagenon A subspecies in a phyletic evolutionary sense, and not a
geographical subspecies.
Wallacea The islands between Wallace’s Line and the boundary of
the Australian Region. Their fauna is of mixed Oriental and Australian
species. (See Map 3.)
Wallace’s Line A line between Bali and Lombok, and points north and
east, which divides the Oriental fauna from that of Wallacea. (See
Map 3.)
Weber’s Line The line of faunal balance between Wallace’s Line and
the boundary of the Australian Region. (See Map 3.)
Weichsel The third and final continental European icecap, correspond-
ing in general to the Wiirm (q.v.) .
Wurm The fourth and last of the Alpine glaciations of the Pleistocene
epoch.
zoogeography The geography of animals; the counterpart to phyto-
geography (q.v.).
zygomatic A paired bone of the face which forms part of the lower and
outer borders of the orbit and parts of its floor. It articulates particu-
larly with the sphenoid temporal and maxillary bones.
ZYGOMATIC
INDEX
Abbevillian industry, 329
ABO blood groups, 173, 193
absorption, concept of, 34
Acheulian industry, 329, 330, 488,
496, 501, 521, 555, 567, 592, 596
achondroplasia, 114-15, 653
acrocentric (telocentric) chromo-
some, 178, 179
Adapidae, 191
adaptation (s) : to crowding, 106—12;
environmental, see environmental
adaptation; general, 28; physio-
logical, see physiological adapta-
tion; social, evolution through,
72-118 passim; special, 28;
unique, of Homo, 118
adaptive threshold, 305
Afalou-Bou-Rhummel remains, 605,
607, 608, 6og, 633
Afghanistan, 330, 482, 484, 526,
559, 577, 578
Africa, 52, 53, 189, 220, 222, 225,
588; apes of, 140, 141, 144-8;
area of, 42; Atlanthropus in, 11;
Bushmen in, see Bushmen; as
cradle of mankind, 656; dwarf
bush bay (loris) in, 113; Eurasia
connected to, 42, 43, 46, 190; and
exchange of animals with South
Asia, 56; fire as late arrival in,
332; Fort Ternan primate in,
205-6; fossil men in, sites of, 590-
609, 612-13; geological events in,
32; human evolution in, 609-10;
land mass of, 42, 43, 46; Lim-
nopithecus in, 196-7, 198; Meso-
pithecus in, 195; monkeys in, 135,
136-8, 139—40; Negroes in, see
Negroes; Pliopithecus in, 198;
pluvial periods in, during Pleisto-
Africa ( continued )
cene, 315; Pygmies in, see Pyg-
mies; tools in, Lower Pleistocene,
227-30, 333; Watusi in, 13, 636;
see also Central Africa, East Af-
rica, North Africa, South Africa,
West Africa
Africanthropus njarasensis, 627
Agache, R., 228, 230
Age of Mammals, 50
agriculture, as ecological grade, 307
Ain Hanech (Hanash), 226, 228,
229, 298
Ainu, 2, 57, 476-7; and facial flat-
ness, 366, 367; hair of, 476; skulls
of, 474, 477; stature of, 456; teeth
of, 355, 357, 516; torus mandibul-
aris of, 451; transportation used
by, 473
Aitape brain case, 399, 406, 410
Alakaluf Indians, 64, 65, 69, 477,
547
Alaska, 318
Alaskan Indians, 63, 65
Aleut language, 5
Algeria, 217, 226, 344, 441, 452,
592, 604, 605, 607
alleles, 36
Allen’s rule, 60
allometry, 25-6, 259
allopatric species, 14
Alouattinae, 132, 133
Alpine race, 19
Alpo-Himalayan system, 189
Alps, 309, 527, 548, 553
amino acids, urinary, in primates,
173
Amphipithecus, 193
anagenesis (succession), 27, 28,
106 ff., 111
Index
ii
Anaptomorphidae, 191, 192
Andaman Islanders, 34, 99, 100,
112, 422, 425
Andamanese language, 407
Andean Indians, 70, 661
anthropoid apes, 140-8
anthropometry, 35
anticlinal vertebrae, and posture,
155
Aotus, 132
apes, anthropoid, 140-8
Arabia, 52, 54, 482, 484, 485
Arabs, 588, 661
Arambourg, Camille, 591, 593, 656
Archaeozoic era, 187 n.
Arctic Circle, 61, 63, 65, 512
Argon-40 dating, 224, 226-7, 313-
14
Asia: Dryopithecus in, 202, 219;
Europe as peninsula of, 42; mon-
keys in, 135-6; see also Central
Asia, South Asia, Southeast Asia,
Western Asia
Assam, 52, 143, 422, 428, 485
Asselar man, 649-50
Ateles, 132, 133
ateliotic dwarfs, 114, 115
Aterian industry, 330, 523, 596, 600
Atlanthropus, 10, 11
Aurignacian industry, 554, 579
Australia, 2, 31, 50, 318, 373; ar-
chaic mammalian fauna in, 92;
area of, 44; arrival of man in, 323,
406; fossil men in, 406—11; land
mass of, 42; monotremes in, 92;
paleolithic tools in, 331; as part of
Australian faunal region, 55; and
placental mammals, 46
Australian aborigines, 2, 4, 5, 59,
91-4, 374, 522; cold-adapted,
66-7, 68; and facial flatness, 367;
hair of, 44; sexual dimorphism in,
26, 27; teeth of, 344, 359, 362,
363; see also Australoids
Australian faunal region, 50, 55, 92,
373
Australoids, 2, 20, 31, 92, 328, 373,
413, 658; cold-adapted, 63, 66,
67; on decline, 660; and facial
flatness, 365, 369; hair of, 44, 426,
427; in India, 18, 373, 422, 485;
languages of, 92, 407; Mongoloids
in contact with, 485, 486; in Ori-
Australoids ( continued)
ental faunal region, 56; teeth of,
352, 353. 426, 453
Australopithecines : cranial measure-
ments and indices of (table), 291;
and Homo, 301-4, 333, 334; teeth
of, 352, 357, 359, 360
Australopithecines: South African,
91, 217, 220, 226, 228, 231-6,
255; arms and hands of, 251-5;
brain case and brain of, 256-60;
cave sites of, 236-7; faces of,
260-4; jaws of, 256, 264-7; legs
and feet of, 244-8; pelvis of,
241-4; postcranial skeletons of,
239-55; shoulder girdle of,
249-51; skulls of, 256, 257-60;
teeth of, 256, 267-77, 359; and
tools, 228, 237-9; vertebrae of,
240, 241
Australopithecus africanus, 232, 233,
236, 243, 255, 259, 270, 271, 272,
274, 276, 277, 290
Australopithecus prometheus, 233
Australopithecus robustus, 236, 243,
255, 259, 267, 270, 271, 272, 274,
276, 290
Austria, 580
autosomal chromosome, 178, 179,
182
axe, hand, paleolithic, 325-7, 329,
330, 484,617
aye-aye, 127, 191
Azande, 432
Azerbaijan, Iranian, 562
Azores, 46
baboons, 137, 139-40, 195
Bakker, E. M. van Zinderen, 644
balanced polymorphism, 22-3
Balangoda skeletons, 424-5
Bali, 55, 113
Bali Strait, 44
Balsequillo, discovery announced at,
479
Baluchistan, 53
Bambandyanolo site, 647
Bantus, 356, 360, 590, 632, 636,
647, 648, 653
Barbary ape, 138, 604
Barbary states, 54
Index
barriers, to movement of land ani-
mals, 46-7, 52, 57
Bartucz, L., 553
Batanta, 55
Bate, Dorothea, 639
Bathurst Island, 94, 103
Bechuanaland, 232, 646, 649
Belgium, 524, 526, 527
Bender, M. A., 179
Benelux, 580
Bengal, 485
Berbers, 54, 588, 604, 609
Bergmann, Carl, 59, 60
Bergmann’s rule, 59, 653
Bering Strait, 43, 54, 69, 190, 314
Besnard, M. V., 649
Bhutan, 18, 143
Biassutti, R., 639
Biberson, P., 595
bilophodontism, 134
binominal, in taxonomy, 9
biogenetic law, propounded by
Haeckel, 164-5
biogeography, 41 n.
Bird, Junius, 478
Birdsell, J. B., 100
Bisitun Cave, 562
Black, Davidson, 431, 432, 434, 437
black apes, 139
Black Earth, Age of, 317
Black Sea, 320, 521, 527, 554, 555,
578
blade tools, paleolithic, 328, 330, 488
Blanc, A. C., 501, 527, 546
blood groups in primates: ABO, 173,
193; MN, 174
Blumenbach, Johann Friedrich, 11
Bodjonegoro, cave at, 413
Boers, 590, 632
Bohlin, Birger, 431
Bolabatu Cave, 415
bolas stones, 228
Bolk, L., 168, 170
Bonch-Osmolovskii, G- A., 555
Bond, W. R. G., 639
Bone, E., 245
Bonobo, 146
Border Cave skull, 632-3
Bordier, F., 228, 230
Borneo, 52, 55, 136, 143, 330, 412,
421
Boskop brain case, 641
Boskop race, 637, 638, 641
iii
Bosporus gateway, 554
Bostanci, E., 561
Boule, M., 12, 410, 437, 543 n., 572,
650
Brace, C. Loring, 510, 554, 657 n.
brachiation, 133, 141, 152, 153, 156,
157, 158, 159, 161, 162
Brachy teles, 133
Brain, C. K., 234
brain: of Australopithecines, 258-60;
of Ganovce specimen, 507; of
Neanderthals, 529; and sapiens-
erectus threshold, 337-46; of
Sinanthropus, 439, 440; size of,
evolutionary increases in, 76-8,
338-41; of Solo man, 394; of Up-
per Paleolithic Europeans, 584;
of Zinjanthropus, 291—2
brain-molar index, 345
brain-palate index, 291-2, 345, 346
Bramapithecus, 203
branching, evolutionary mechanism
of, 27, 28
breeding areas, for land animals, 46,
47, 49
Breuil, H., 501-2, 627
Briggs, L. C., 604, 607, 632, 648
British Isles: Cresswellian industry
in, 579; Neanderthals in, 525; and
Palearctic faunal region, 54;
Swanscombe skull from, 91, 314,
487, 495-7; Upper Paleolithic fos-
sil man sites in, 580
Brodar, S., 508
Broken Hill man, 336, 337, 341, 344,
346, 442, 621-7; cranial capacity
of, 623; face of, 625-6; teeth of,
624-5
Bronze Age, 108, 423
Broom, R., 231, 245, 246, 252, 253,
267, 300, 636
Brothwell, D., 566
Bruckner, E., 310, 311, 312, 314
Buginese, 415
Burma, 53, 143, 193, 330, 422
Bushmen, 4, 27, 57, 59, 63, 68, 99,
588, 589, 590, 630, 632, 636, 637,
640, 641, 646, 660; and facial
flatness, 364, 366, 367; Kung, 100,
102; number of, 660; pedomor-
phism in, 646; rock paintings by,
638, 649; teeth of, 344, 353, 354,
359. 360, 362, 364, 455, 456; and
IV
Index
Bushmen ( continued )
Temefine-Tangier line, 601, 602,
649
Bushnell, G., 521, 522
Butler, P. M., 211, 212, 214
California: early glacial till of, 314;
Indians of, 34, 99, 546
Callenfels, P. van Stein, 413, 414,
420
Callithricidae, 131
calva, defined, 256
calvarium, defined, 256
Cameroons, 147, 651
Canary Islands, 604
cannibalism, 432, 601, 602
Canoe Indians, of Tierra del Fuego,
64
Cape Flats skull, 630-2
Cape Martin, Australia, 406, 407
Capoids, 2, 4, 57, 328, 590, 630,
654, 656; and facial flatness, 366,
369; modern, formation of, 645-9;
in North Africa, 485; origin of,
636-9; shrinking process in, 645,
646; teeth of, 354, 360, 364
Capsian culture: in East Africa, 607,
634-6; in North Africa, 330, 606-7
Capsian remains, 278, 609, 634, 635,
636
capuchins, 132
Carabelli’s cusp, 363, 516, 635
Caribs, 432
Carpathian Mountains, 554
carrying angle, in locomotion, 157,
158, 160
Caspian Sea, 42, 320
catarrhines, 131, 132, 133, 134;
evolution of, 192-6
Caucasoids, 2, 12, 27, 66, 328, 658,
660, 663; contacts of, with other
subspecies, 485-6; European, see
European Caucasoids; and facial
flatness, 364, 365, 367, 369; geo-
graphical distribution of, 59;
homeland of, search for, 482, 484;
in India, 18, 59, 374, 422, 482,
484; merged into Mongoloids, 18;
in North Africa, 52, 482, 588, 590,
603; in Oriental faunal region, 56;
and physiological adaptation to
climate, 63, 69; Tasmanians ab-
Caucasoids ( continued )
sorbed by, 34; teeth of, 352-5,
360-4; in Western Asia, 482, 484,
485, 487, 498; see also Neander-
thals, Upper Paleolithic Euro-
peans
Caucasus Mountains, 12, 554
Cave of Hearths mandible, 628-9
cebid monkeys, 131-3
Ceboidea, 193
Cebus, 132
Celebes, 55, 56, 112, 113, 139, 414,
415
Cenozoic era, 50, 187-8, 190, 216
Central Africa: Pygmies in, see Pyg-
mies; Sangoan industry in, 330;
Watusi in, 13
Central America, 54
Central Asia, dines in, 18; Soviet,
482, 526
Central European Neanderthals,
549-54; mandibles of, 550-2;
postcranial bones of, 552; and
Rumanian toe bone, 553; signifi-
cance of, 553-4; and Subalyuk
child’s skeleton, 552-3
Central Honshu remains, 471-2
centromere, 178
Ceram, 56
Cercocebus, 137
Cercopithecidae, 133-5, 141> 194>
198, 214
Cercopithecinae, 135, 136-40, 162,
195
Cercopithecoidea, 193
Cercopithecus, 137
cerebral evolution, level of, 340
Ceylon, 52, 99, 139, 366, 367, 414,
422, 423, 424, 425, 518
Chancelade man, 577, 583, 584-5
Changyang maxilla, 461-2
Chao, T-K., 449
Chardin, Teilhard de, 431, 470
Chatham Islands, 62
Cheirogaleus, 127
Chellian industry, 329
Chellian-3 skull, 336, 337, 614,
616-17
chest, and locomotion, 162, 167
Chia, L-P., 461, 476
chimpanzees, 140, 144-6; ancestors
of, 198 ff pygmy, 146
chin, rise of, 346-50
Index
v
China, 8, 205, 222, 223, 225, 230,
302; and Australopithecines,
300-1; Dryopithecus teeth from,
203; early skeletal material from
(table), 430; fossil macaques in,
195; geological regions of, 316-17;
Miao tribes in, 416-17; monkeys
in, 136, 139; and Oriental faunal
region, 52, 53; paleolithic tools in,
331; Pleistocene apes in, 206-7;
possible survivals of apes in,
207-8
Chinese: cold-adapted, 65, 362;
torus mandibularis of, 451
Chingshui Erosion, 317
choppers and chopping tools, paleo-
lithic, 228, 325, 326, 328, 330,
33i>478
Choukoutien, 91, 103, 302, 323, 337,
43°. 431, 434, 436, 461; geology
of, 435; pollen analysis of breccia
from, 436; Upper Cave of, 337,
472-5
chromosomes, primate, table of,
180-2; taxonomy aided by study
of, 177-83
cingulum, defined, 357
Cipriani, L., 648
Clactonian industry, 329, 500, 521
Clark, Desmond, 237, 332, 622
Clark, LeGros, 219, 243, 388
cline, defined, 18
Clovis industry, 479
Cohuna skull, 409-10
Colobinae, 133, 135-6, 195
Congo, 113, 135, 147, 589
Congoids, 2, 4, 328, 485, 588, 633,
649, 656, 658
Coolidge, H. J., Jr., 146
Coppen, Yves, 297
cranium, defined, 256
Cresswellian industry, 579
Cretaceous period, 188 n., 190
Crimea, 505, 526, 555
Crocuta crocuta, 321, 435, 489
Cro-Magnon man, 35, 346, 472, 577,
582
Cromerian Interglacial, 222, 223,
229,310,316, 331,435
crowding: adaptation to, 106-12;
and domestication, 117; dwarfing
as solution to problem of, 112-15
Curtis, G. H., 226, 227, 616
Cynopithecus, 139
Cyphanthropus rhodesiensis Wood-
ward, 626
cytogenetics, experimental, 182
Czechoslovakia: Ganovce stone
brain from, 487, 507; Mesopithe-
cus in, 194; Neanderthals in, 525,
527, 55°; Upper Paleolithic fossil
man sites in, 581, 582
Danger Cave seed-gathering site,
479
Darling Downs, South Queensland,
408
Darlington, P. J., 52, 53
Dart, R. A., 231, 232, 233, 235, 238,
239» 242, 628, 632, 656
Darwin, Charles, 12, 50, 111, 116,
151, 164, 656
Dasht-i-Lut desert, 189
dating: Argon-40, 224, 226-7,
313-14; Carbon-14, 310> 313;
with sea-water isotope method,
312
Daubentonia, 127
Davies, P. R., 201, 202
Dead Sea, 222
Dendrogale, 121
Deraniyagala, Paul, 424, 425
dimorphism, sexual, 26-7
Dinaric race, 19, 35
diploid cell, 177
Dire Dawa mandible, 627-8
Djetis faunal beds, 223, 224, 225,
229, 298-9, 314, 316, 323, 331,
344. 375, 383
Doherty, J. G., 575
domestication, biological results of,
116-18
dominance: of groups, 48-50, 661; as
resistance to intrusion of outsiders,
661
Donau glaciations, 222, 314
Dong Thuoc skull, 419
Dordogne region, France, 579
Drennan, M. L., 631, 632
Dreyer, T. F., 642, 644
drills, West African, 140
Dryopithecinae, 141, 199, 202, 203,
204, 218, 219, 223, 334, 360
Dryopithecus pattern, 141, 360, 361,
363
VI
Index
Dubois, Eugene, 384, 386, 401,
410,413
DuBrul, E. L., 74, 347
Duckworth, W. L. H., 421
Duwwuds, 648
dwarfing, 34, 426; achondroplastic,
114-15; ateliotic, 114, 115; as
solution to problem of crowding,
112-15; see also Pygmies
East Africa, 57, 91, 225, 228, 590,
617, 653; Capsian culture in, 607,
634-6; early hominids in, 277-97,
304; hand-axe period in, fireless,
332; Kanjeran Pluvial period iden-
tified in, 315; lakes of, 321;
Levalloisio-Mousterian industry in,
330; Lower Pleistocene tools in,
227; Mesopithecus in, 194; patas
monkey in, 137
ecology, 58, 150
Egbert, from Ksar ‘Akil site, 575
Egypt. 193. 639
Egyptians, ancient, 366, 367
Ehringsdorf remains, 487, 505-6,
512-13,514
Elementeita series, 634, 635
Elster glaciation, 310
embryo, 23, 49, 164, 166
embryology, 164-71
Emiliani, Cesare, 312, 313, 314
enamel pearl, 357, 359
endocrine system, 24, 109; and tem-
perament, 115-16
England, see British Isles
environmental adaptation, and early
man, 58-9; evolution through,
38-71
Eoanthropus, 437
Eocene epoch, 32, 189, 190, 191,
218
Equas stenonis, 229
erect posture, 149, 156, 160, 161,
168; carrying angle as adapta-
tion to, 157; and teeth, relation-
ship to, 153-4, 162-4
erectus-sapiens threshold, see sapi-
ens-erectus threshold
Erythrocebus, 137
Eskimo, 2, 62; faces warmed by
extra blood flow, 61, 65, 534; and
facial flatness, 367, 369; language
Eskimo ( continued )
of, 5; stature of, 456; teeth of,
355. 357, 360, 362, 363, 455, 456,
516; toms mandibularis of, 451
Ethiopia, 135, 140, 627
Ethiopian faunal region, 50, 52, 53,
54, 56, 57, 92, 189, 485
Eurasia: Africa connected to, 42, 43,
46, 190; area of, 42; land mass of,
42, 46; Lower Pleistocene tools in,
228; and Palearctic faunal region,
54
Europe: arrival of man in, 57; Dryo-
pithecus in, 202, 219; fossil men
in, 487-8, 497 ff.; gibbons in, an-
cestral, 197, 198; macaque
fossils in, 195; Neanderthals in,
488; Palearctic genera in, 57, 321;
paleolithic tools in, 11, 329, 330;
as peninsula of Asia, 42; glacia-
tions and interglacial ages in, 222,
309 ff., 486, 487; recession of last
ice sheets in, 189, 486
European Caucasoids, 2; and phy-
siological adaptation to climate,
63, 69; sexual dimorphism in, 26;
teeth of, 354, 361, 362
euryphagous species, 15
Evans, I., 421
Evernden, J. F., 226, 227, 616
evolution: body size as factor in,
38-9; and brain size, 76-8, 338-
41; through branching, 27, 28; of
catarrhines, 192-6; climate as fac-
tor in, 40; earth’s face as determi-
nant of, 41-2; through environ-
mental adaptation, 38-71; and
fossil primates, 215-16; isolating
mechanisms in, 103-6; and laws of
change, 8; mutation as primary
element in, 21; parallel, 11, 37,
132, 192; phvletic (succession),
27, 28, 106 ff., 111; of platyr-
rhines, 192; through social adap-
tation, 72-118; space requirement
as factor in, 39; and species for-
mation in, 21-2; through succes-
sion, 27, 28, 106 ff., 111
Ewer, R. F., 234
Ewing, J. Franklin, 575
Eyasi man, 627
eye color gradient, 18, 149
Index
Vll
face(s): of Ainu, 366, 367; of Aus-
traloids, 365, 369; of Australo-
pithecines, 260-4; of Broken Hill
man, 625-6; of Bushmen, 364,
366, 367; of Capoids, 366, 369; of
Caucasoids, 364, 365, 367, 369,
446; of Indians, American, 369,
474; of Mongoloids, 364, 365,
366, 367, 369, 428; of Neander-
thals, 534-5; of Negritos, 369; of
Negroes, 366, 367, 369; of Pyg-
mies, 654; of Sinanthropus, 445-7;
of Solo man, 396-7; of Ternefine
man, 595; of Upper Paleolithic
Europeans, 584; of Wadjak man,
403, 427, 445; of Zinjanthropus,
289
facial flatness, as criterion of race,
364-9; indices of, 367
Fairbridge, Rhodes W., 312, 313,
314,318, 477
Fairservis, Walter A., Jr., 393
faunal association, 234, 235
faunal balance, line of, 55
faunal regions, 50, 52-5; and human
origins and movements, 56-8
Fayum, 193, 196
Fen Valley flints, 522
Ferenbach, Denise, 605, 607
Fergusson Island, 426
fetus, 23, 49, 164, 165, 167, 168,
170
Fiedler, W., 120, ig8
Fiji Islands, lg
Finns, 68, 515
fire: chronology and distribution of
use of, 332; discovery of, 90-1;
evidences of, 91, 229, 302, 332
First Himalayan Glaciation, 310
First Wiirm Interstadial, 329, 330
Fish Hoek skeleton, 645
Fitzsimmons, F. W., 641
flake tools, 228, 327, 329, 330, 331,
5°i
Florisbad cranial fragment, 638,
642-5
Flower, H. W., 352
Flower’s index, 353, 426
Folsom site, 478, 479
Fontechevade skulls, 487, 498-500
food gatherers, living, 91, 288; fossil
men compared with, 99; mating
food gatherers (continued)
systems among, 102; population
size among, 100-2
foot of primate, in fetal life, 168
Formosa, 52, 139
Formosov, A. A., 557
Fort Ternan primate, 205, 206, 209,
215, 218, 220, 287, 334
fossil men, 8, 10, 25, 79; in Africa,
sites of, 590-609, 612-13; in Aus-
tralia, 406-11; genetics applied to
study of, 35; grades and species
°f, 332-7; fines and subspecies of,
350 ff.; living food gatherers com-
pared with, gg; longevity of, 103;
in North Africa, sites of, 590-609;
racial differences among, 63; of
Riss-Wiirm Interglacial age, 487,
49 7> 498; sexual dimorphism in,
26; smallest cranial capacity of,
260; temporal and spatial distribu-
tion of, 322-4; tools of, see tool-
making; in western Asia, 487, 498,
587; see also Upper Paleolithic
Europeans
fossil record, 186-216; and human
evolution, 215-16
France: Fontechevade skulls from,
487, 498-500; Lower Pleistocene
deposit in, 228—9; Mesopithecus
in, ig5; Monsempron remains
from, 487, 511, 512, 514, 515;
Montmaurin remains from, 487,
511, 512, 514, 515; Neanderthals
in, 524-5, 526, 527; paleolithic
archaeology born in, 324; Perigor-
dian industry in, 579; Upper
Paleolithic fossil man sites in,
580, 582
Fuegians, 31, 34, 64, 68, 457, 480,
546
Galago elegantulus, 129
galagos, 129
Galapagos Islands, 50
Galilee skull, 566, 567
Galloway, A., 647
Gamble’s Cave, 634, 635, 636
Gamblian Pluvial period, 315, 622,
634
gammaglobulin (Gm) test, 175
Ganovce stone brain, 487, 507
Index
viii
Garos, 422
Garusi site, early hominids found in,
277> 295
Gates, R. R., 654
gelada, 137, 140
general adaptation, 28
genetic drift, 47-8
genetics, 12, 14 n., 21, 22-4, 35, 36;
status quo in, 662; taxonomy
aided by study of, 177-83
Gentner, W., 224
genus, in taxonomy, 9
geological time, divisions of, 187-8 n.
Gerasimov, M. M., 557, 559
Germany: Ehringsdorf remains from,
487, 505-6, 512-13. 514; Mauer
mandible from, 91, 347, 382, 449,
452, 487. 489-92, 593; Neander-
thals in, 524, 526, 527; Paido-
pithex in, 203; Steinheim cranium
from, 341, 487, 492-5; Upper
Paleolithic fossil man sites in, 580,
582
gerontomorphism, 25
gibbons, 140, 141, 142-3; evolution
of, 196-8
Gibraltar, 138, 525
Gibraltar, Strait of, 42, 320
Gigantopithecus hlacki, 206-7, 219,
300
glacial geography, 318-22
Gleiser, I., 359
Goodwin, A. J. H., 638
gorillas, 140, 141, 147-8; ancestors
of, 198 ff .
Gorjanovic-Krambergcr, K., 508,
5io,5i3,5i6
Gottweig Interstadial, 330, 390, 412,
486, 488, 531, 549, 554, 562, 577,
578, 579, 602, 603
Grabham, G. W., 639
grades, ecological, concept of, 305,
306, 334
Great (Mindel-Riss) Interglacial
age, 310, 311, 314, 329, 442, 486,
488, 492, 496, 498, 521
Greenland, 54, 70, 92
Gregory, W. K., 151, 220 n., 360,
389
Gremiatskii, M. A., 561
Grimaldis, Negroid, 577, 583, 584
growth, postnatal, differences in,
171-2
Guak Kepah, mandible from, 420
Guam, 445
Giinz glaciations, 222, 309, 310, 312,
313,314,315, 435
Giinz-Mindel Interglacial age, 310
Gusinde, M., 654
Haeckel, Ernst, 164, 165
hair: of Australoids, 44, 426, 427;
color of, 149; fetal, 168, 170; of
Mongoloids, 428; of Pygmies,
654
Halmahera, 56
Hamites, 637, 638, 646, 654, 656
Hammel, H. T., 64, 66
hand ax, paleolithic, 325, 326, 327,
329, 330, 484, 617
hands, cold-adapted, 65; uses of,
154
haploid cell, 177
haptoglobins, in primates, 175
Harrison, Tom, 412
Hart, C. W. M., 97
Haua Fteah mandible, 602-3
Heberer, K. A., 430
Heck, H., 146
Heidelberg jaw (Mauer mandible),
9i, 347, 382, 449, 452, 487,
489-92, 593
Heilman, Milo, 220 n., 389
hemoglobins, in primates, 175
Henri-Martin, G., 498
Henry, Jules, 110
Higgs, Z. S., 566
Himalayas, 18, 46, 70, 136, 138, 310
Hindu Kush Mountains, 520, 526,
554
Ho tribe, 420, 422
Hoffman, A. C., 644
Hokkaido, 330, 474
Holarctic faunal region, 52
holotype, in taxonomy, 1 1
Homa shell-mound skulls, 640-1
home range of animal, 81
Hominidae, 148-50; distinguished
from Hominoids, 208—9; distribu-
tion of early, 231; divisions of,
220; earliest, 217-304; geography
and numbers of early, 230 ff.; tool-
making as behavioral character-
istic of, 227
Index
IX
Hominoids, 193; distinguished from
hominids, 208—9; earliest known
specimens of, 196 ff.
Homo erectus, 24, 119, 323, 384,
385* 399, 477, 482; absence of
chin in, 349; and Neanderthals,
530; oldest known, 33; physiologi-
cal adaptation by, 69; and
sapiens-erectus threshold, see
sapiens-erectus threshold; skulls
of, 278, 292, 341, 343, 494; tooth
size of, 344, 345; transition to
Homo sapiens, 16, 27, 30, 33, 39,
in, 3°6, 374, 427, 481
Homo modjokertensis, 375, 383-4,
440; see also Pithecanthropus
modjokertensis
Homo rhodesiensis, see Broken Hill
man
Homo sapiens, 5, 9, 10-11, 39, 119,
323; chin of, 346; cranial form of,
341, 3435 as dominant group, 49;
evolutionary changes within,
346-50; first appearance of, 33;
grades of, 336; pedomorphism in,
25, 161; polymorphism in, 17,
161; ratio between brain size and
palate area, 292; and sapiens-erec-
tus threshold, see sapiens-erectus
threshold; sexual behavior of,
83-4; tested for physiological ad-
aptations, 63, 70; tooth size of,
344, 345; transition from Homo
erectus, 16, 27, 30, 33, 111, 306,
374, 427, 481
Hong Kong, 52
Honshu remains, 471-2
Hooijer, D. A., 223
hoolock, 143
Horton, W. E., 632
Hottentots, 4, 367, 589, 630, 633,
637, 645, 646, 647, 649, 660
Hotu cave, 587
Howell, F. Clark, 223, 236, 237,
304, 395, 49i,493,6io
howler monkeys, 132, 133
Hrdlicka, A., 566, 622
Hsiin-Tzu, 207
Hungary: Neanderthals in, 525,
527, 549, 55i; Upper Paleolithic
fossil man sites in, 581; Solutrean
industry in, 579
Huns, 484
Hunt, C. B., 227
Hunt, E. E., 359
hunting: beginning of, 79-80; as
ecological grade, 307; speech
necessary for, 80, 87
Hiirzeler, J., 210, 212
Huxley, Thomas, 151
hybridization, 12
Hylobates, 142
Hylobatidae, see gibbons
Hylobatinae, 141
hypothalamus, 109
Ice Age, 310, 311
Illinoisan glaciation, 310, 312, 313,
314, 318, 477, 478
index (indices): brain-molar, 345;
brain-palate, 291-2, 345, 346;
cranial, 668 ff.; of facial flatness,
365-6, 367; Flower’s, 353, 426; of
mandibles, 450, 675
India, 2, 20, 54, 222, 328, 329, 366,
367, 373, 407; Australoids in, 18,
373, 422, 485; as breeding place
of Hominidae, 205; Caucasoids in,
18, 59, 374, 422, 482, 484; clinal
zone in, 18; Dryopithecine jaws
and teeth from, 203, 218-19;
Kadars in, 52, 100; macaques in,
138, 139; Mesopithecus in, 195;
Mongoloids in, 422; and Oriental
faunal region, 52; prehistoric pop-
ulations of, 422-3; pre-Soan indus-
try in, 229; tribal peoples of, 2,
18, 422
Indians, American, 2, 307, 445, 474,
4 77, 478, 479, 480; and facial flat-
ness, 369; as Mongoloids, 477,
479, 480; nose form of, 428; teeth
of, 355, 356, 455
Indochina, 44, 52, 53, 143, 195, 330,
422, 426; Mesolithic and Neo-
lithic remains from, 416-21
Indonesia, 20, 34, 44, 52, 56, 92,
121, 139, 318, 330, 373, 422, 425;
Mesolithic-Neolithic transition in,
413-16
Indus Valley, 423
insectivores, 121
interbreeding of species, 12-13
interfertile species, 12
interstadial, defined, 329 n.
X
Index
I
(
]
1
I
/
<
I
f
Iran, 194, 482, 526, 562, 577, 578,
581, 587
Iraq, 484, 526, 562, 577
Iron Age, 34, 108, 636
ischial callosities, 134-5, !39> 141 ,
142, 145, 162
ischium, in locomotion, 159, 160
isolating mechanisms, in evolution,
103-6
isolation, defined, 22
Israel, 229, 237, 297-8, 482
Italy, 22i, 320, 522, 523; fossil
macaques in, 195; Late Acheulian
site in, 314; Neanderthals in, 525,
526, 527; Pontian fossil beds in,
209; Saccopastore remains from,
487, 500-4, 514, 515, 522, 527;
Upper Paleolithic fossil man sites
in, 580
Japan: Ainu in, see Ainu; dwarf
animals in, 112, 113; early skeletal
material from, 429, 430, 460,
464-5, 471-2, 476; macaques in,
139
Japanese, 31, 65, 67, 456, 517
Java, 4, 52, 91, 143, 217, 220, 222,
301> 3°4> 3i5> 33°, 332, 410, 413;
Djetis faunal beds in, 223, 224,
225, 229, 298-9, 314, 316, 323,
33i, 344, 375, 383; Meganthropus
mandibles from, 298-300, 301;
Solo skulls from, see Solo skulls;
Trinil fauna in, 224, 260, 314,
316, 337, 384-6, 387 ff-i Wadjak
man in, see Wadjak man
Jebel Qafza remains, 566, 567
Jerison, H. J., 340
Jolly, Keith, 619
Jones, F. Wood, 129
Jordan Valley, 79, 80, 222, 229, 239,
297
Jurassic period, 188 n., 189
Kabuh beds, 316
Kadars, 52, 100, 102
Kagerian Pluvial, 315
Kaiso fauna, 225, 228
Kait’o-Tung Cave specimen, 475-6
Kalahari Desert, 68, 100, 590, 646
Kalin, J., 214
Kamasian Pluvial, 315
Kanam site, 225, 235; mandible
from, 295-7
Kanjera specimens, 617—19
Kanjeran-Gamblian Interpluvial, 323
Kanjeran Pluvial, 315, 617
Kansan glaciation, 310, 312, 313
Kartan culture, 406
karyotype chart, 179, 182
Kashmiri, 661
Kedung Brubus, 384, 385
Kei island, 55
Keilor skull, 407-8, 410
Keith, Arthur, 292 n., 345 n., 566,
57U 572, 576
Kenya, 195, 196, 205, 219, 277, 296,
332, 618, 634, 635, 640
Keo Phay skull, 420
Khartum Negroid specimen, 651
Khasis, 52, 422
Khyber pass, 53
Kiik-Koba cave, 555-6
Kinsey, A. C., 83
Kirghiz, 18
kladogenesis (branching), 27, 28
Klatt, Berthold, 117
Kohl-Larsen, Ludwig, 295, 627
Kokten, I. K., 561
Korana, 630, 637
Kourounkorokale, 650
Krapina remains, 487, 508-11,
513-14, 515, 516-19
Kromdraai site, 232, 233, 234, 235,
236, 238; elbow from, 251-2;
metacarpal from, 254; skull frag-
ment from, 258, 266; talus from,
246-7
Krzywicki, L., 100, 101
Ksar ‘Akil site, 575
Kubu tribe, 421
Kummer, B., 170
Kung Bushmen, 100, 102
Kurten, B., 10, 222 n., 435
Kwangsi province, China, 317, 467,
475
La Chapelle aux Saints, 12, 520,
528; clavicle of, 543; cranium of,
531; crippled by arthritis, 103,
542, 543; feet of, 547; femora
of, 546; hands of, 545; height of,
548; humeri of, 544; kneecaps of.
Index
xi
La Chapelle aux Saints ( continued )
546; mandible of, 535, 537; nose
°f, 532, 533; pelvic bone of,
545-6; postcranial skeleton of,
542, 543; ribs of, 543; teeth
of, 539, 542; tibia of, 546; ulnae
of, 544; vertebral column of,
542-3; wrist bones of, 545
Laetolil fauna, 225, 228, 295
Lagothrix, 133
Lamontjong Cave, 415
land masses, 42—4, 46
Lang Cuom skulls, 419, 420
Langhnaj skeletons, 422-3
language (s), 86, 107; Australian,
92, 407; diversity of, 5; Indian,
479; Mon-Khmer, of southeast
Asia, 407, 422; nontonal, 5; tonal,
5; see also speech
langur, 135, 136, 138
Laos, 416, 417
Lapland, 70, 92
Lapps, 63, 65, 66, 67, 68, 359, 451,
646
Lartet, Edouard, 202
Last Interglacial, see Riss-Wiirm
Interglacial
Last Interstadial, 497
Late Acheulian site, in Pietra, Italy,
314
leadership, in human relations, 72-3,
86, 108—9
Leakey, L. S. B., 80, 202, 205, 219,
227, 238, 278, 284, 285, 287, 288,
289, 291, 332, 610, 614, 616, 617,
634. 636, 641, 656
Lebanon, 330, 484, 526, 555, 575,
577, 578
Lehner mammoth site, 478
lemurs, 120, 126-8
Lesser Sundas, 55
Levalloisian industry, 329, 521
Levalloisio-Mousterian industry,
329, 330, 501, 523, 603
Levant, 320, 555
level of cerebral evolution, 340
Lewis, G. E., 203, 219
Li, Y. H., 207, 239
Liang Annals, 207
Liberia, 113, 651
Libya, 591, 602, 648
lice, mutual, on primates, 176, 193
Licent, E., 470
Limnopithecus, 196, 197, 198
Limpopo River, 589, 647, 648
line(s) : concept of, 305, 306, 308;
of fossil men, 350 ff.; gibbon,
196-8; Negro evolutionary, 611,
613-14; Pithecanthropus, 373-90;
Pithecanthropus- Australoid, 410-
11, 427; Ternefine-Tangier, 600-2,
649, 658
Linnaeus, 9, 13
Lippolt, H. J., 224, 226, 227
Litorina Cave mandible, 595
Liu-Kiang man, 467-70
Livingstone, Frank, 657 n.
local races, defined, 19
Lombok, 55
Lompoa Cave, 415
Loris tardigradus, 129
lorises, 120, 128-9
Loth, E„ 537
Lower Pleistocene epoch, 57, 118,
i95> 207, 217, 220, 221-6, 305,
310, 313, 320, 321, 660; African
archaeological sites in, 227, 590;
Australopithecines and Homo
during, 301, 333; Djetis fauna in,
316, 323; new dating for, 226-7;
of South China, 316-17; tool-
making in, 227-30, 333
Luzon, 330
macaques, 137, 138-9, 195
Macassar Strait, 44
McBurney, Charles, 521, 522, 602,
604, 651
McCown, T. D., 571, 572, 576
Madagascar, 52, 126, 128, 189,
428, 660
Magdalenian industry, 579, 582
Makapansgat site, 232-9 passim,
628; brain case from, 258; clavicle
from, 249; femoral head from,
245; humerus fragment from, 251,
252; pelvic bones from, 241, 243;
teeth from, 270
Malaya, 44, 52, 112, 143, 228, 229-
30, 330,414, 421
Malez, M., 554
mammoth, hunting of, 101, 150
mammoth site, Lehner, 478
Manchuria, 65
mandible, defined, 256
xii Index
mandrills, 140
Mapa skull, 411-12, 462-4, 477
Mapungubwe site, 647
Marks, P., 299
marmosets, 131, 132
Marrett, J. R. de la H., 424
marsupials, g2
Martin, Rudolf, 516
mating systems, 81-2; among food
gatherers, 102
Matjes River skeletons, 645
Matthew, W. D., 50
Mauer mandible (Heidelberg jaw),
9b 347, 382, 449, 452, 487,
489-92, 593
Mauritius Island, 50
maxilla, defined, 256
Mayr, E., 437
Mediterranean race, 19
megadont teeth, 353
Meganthropus, 299, 301, 375, 382
meiosis, 21
Melanesia, 44, 373, 489
Melanesians, 2, 112, 353, 367, 369,
420, 425, 517
melanin, 69
Melville Island, 94, 331, 346
Mendel’s second law, 21
mesodont teeth, 353
Mesolithic remains, in Ceylon, 424;
in Indochina, 416; in Indonesia,
413; in North Africa, 605, 606-9
Mesopithecus, 194, 195
Mesozoic era, 187 n., 189
metabolism, purine, 172, 193
metacentric chromosome, 178, 179
Mettler, L. E., 179
Mexico, 54, 190, 479
Miao tribes, 416-17
microdont teeth, 353
microliths, 325, 328, 330, 331, 509
Micronesians, 2
Middle Eastern whites, 2
Middle Pleistocene epoch, 57, 217,
220, 223, 305, 321, 332, 334, 484,
630, 658, 660; beginning of, 221,
222, 225, 230, 301, 302, 304, 310,
313, 314, 425; deposits of, in
north China, 317; fossil man sites
in, 322, 323, 374-5, 460-4, 591,
592; hunting in, 80; tool types in,
328, 329, 333; Trinil fauna in, 316
Mijsberg, W. A., 414, 415, 420-1
Milankovitch, M., 311, 313, 314
Miles, J. R. E., 211, 212, 214
Mindanao, 55
Mindel glaciation, 223, 309-15 pas-
sim, 476
Mindel-Elster glaciation, 435
Mindel-Riss (Great) Interglacial,
310, 311, 314, 329, 442, 486, 488,
492, 496, 498, 521, 578
Miocene epoch, 32, 189, igo, 216,
218, 220, 283; Dryopithecus in,
202, 219; and Italian fossil beds,
209; Limnopithecus in, 196-7,
198; Mesopithecus in, 194; Plio-
pithecus in, 197, 198
Misol, 55
Mivart, St. George Jackson, 151
MN blood groups, 174
Molucca Islands, 55, 406
Mongoloids, 2, 20, 26, 27, 31, 56,
328, 374, 407, 413, 417, 421, 422,
658, 660, 663; Australoids in con-
tact with, 485, 486; Bering Strait
“highway” crossed by, 477, 478,
47g, 480; Caucasoids merged
into, 18; cold-adapted, 63, 65, 66,
69; and facial flatness, 364, 365,
366, 367, 369, 428; geographical
distribution of, 59; hair of, 428; in
high plateaus, 70; as homogeneous
subspecies, 428; in India, 422;
languages spoken by, 5, 407; nose
form of, 428; scapular spine of,
517; skin color of, 428; teeth of,
352, 353, 355, 356, 357, 359,
360, 362, 363, 364, 428, 453
Mongols, 65, 484
monkeys, New World, 131, 132,
133, !93; Old World, 131, 132,
!33> 134-40, 141, 193
Mon-Khmer languages, 407, 422
Monod, T., 649
monotremes, 92
monotypic species, 14
Monsempron remains, 487, 511, 512,
514, 515
Montmaurin remains, 487, 511, 512,
514, 515
Morant, G. M., 5, 8, 365, 527, 582
Morocco, 2, 4, 138, 301, 344, 591,
595, 605, 607
mosaic, defined, 18
Mouillian culture, 330, 604-5
Index
xm
Mount Carmel postcranial skeletons,
572-5, 576
Mount Carmel teeth, 571-2
Mousterian industry, 329, 501, 505,
508, 512, 520, 522, 523, 527, 549,
550, 55i, 554, 555, 558, 562
Movius, H. L., Jr., 460, 523, 559,
579
Munda tribe, 420, 422
Murdock, G. P., 102
Mussolini cave, see Subalyuk cave
mutation: defined, 21; favorable,
chances for, 47
Nagas, 423, 428
Naivasha Railroad Station remains,
634, 636
Nakuru burial ground, 636
Napier, J. R., 201, 202
Nasalis, 136
National Geographic Society, 226
natural selection, Darwin’s theory of,
12, 111
Neander, Joachim, 519
Neanderthals, 5, 8, 12, 13, 35, 61,
323, 336, 342, 346, 455, 456,
464, 488, 519 ff.; Central Euro-
pean, see Central European Nean-
derthals; dead buried by, 539;
longevity of, 103; numbers and
distribution of, 523-7; origins of,
575-7; Eastern, of Shanidar,
561-5; from Soviet Union, see So-
viet Union Neanderthal remains;
Western European, see Western
European Neanderthals
Neanthropic grade, 334, 336
Nearetic faunal region, 50, 52, 54,
57, 149, 221
Nebraskan glaciation, 310, 312, 313
Negritos, 2, 413, 425; and facial flat-
ness, 369; as food gatherers, gg;
in Oriental faunal region, 56;
Philippine, 112; of Tam Hang,
417-18; teeth of, 353
Negroes, 2, 22, 26, 112, 588, 589,
658; and facial flatness, 366, 367,
369; heat adapted, 63, 68-9; mod-
ern, earliest skeletons of, 649-51;
and possible evolutionary line,
611, 613-14; and Pygmies, gene
flow between, 652, 653; scapular
Negroes ( continued )
spine of, 517; sickling trait carried
by, 22, 70; teeth of, 353-4, 356,
360, 361, 362, 363; theory of
origin of, 655-6; urbanized, high
blood pressure in, 110; vitality of,
660
Negroid, defined, 4
Neolithic era, 34, 102, 413, 416,
420, 587, 650
neoteny, 24
Neotropical faunal region, 50, 54,
141, 189
Nepal, 18
Nepalese, 366, 367
Neumann, G., 480
New Caledonia, 30-1, 421, 425, 426,
489
New Guinea, 44, 55, 56, 92, 309,
373, 406, 410, 426
New World monkeys, 131, 132, 133,
193
Ngandong leg bones, see Solo leg
bones
Ngandong skulls, see Solo skulls
Niah Cave, Upper Pleistocene skull
from, 411, 412, 413
Nihowan-Taiku beds, 316
Nile Valley, 635
Nordic race, 19, 35
North Africa, 226, 315, 330, 523,
589> 59°, 637, 660; Atlanthropus
in, 11; Barbary ape in, 138, 604;
Capsian skeletal material in, 606-
7; as Caucasoid territory, 52, 482,
588, 590, 603; Ethiopian fauna of,
52; fossil men in, sites of, 590-
609; Lower Pleistocene tools in,
227, 228; Mesolithic remains in,
racial anatomy of, 607-9; Mouil-
lian invasion of, 330, 603-5; pre-
Mouillian skeletal material from,
591-603
North America, 54, 477; area of, 43;
arrival of man in, 477, 478, 479,
480; industries of tools in, 331,
478, 479; land mass of, 42, 43,
46; recession of last ice sheets in,
189; South America connected to,
43; 190
Northeast Frontier Agency, 18
Northern Hemisphere, 46, 57, 189,
3ii
XIV
Index
!
:
nose form, and climate, 62, 533-4
Notopuro beds, 316, 390
Nubians, 366, 367
Oakley, K. P., 236
Oceania, 2, 4, 56, 112, 460
Ochoz mandible, 550, 551
Okladnikov, A. P., 558
Old World monkeys, 131, 132, 133,
1-34-4°, Mi. *93
Oldowan tool level, 234, 235, 278,
281, 301
Olduvai child, 80, 247, 278, 281,
334; clavicle of, 285; evolutionary
and taxonomic position of, 287;
finger bones of, 285, 286; foot
bones accompanying, 284-5;
hands of, 286; mandible of, 281-2;
parietal bones of, 283-4; teeth of,
282-3
Olduvai Gorge, 225, 228, 229, 237,
239, 298, 316, 325; Bed I of, 79,
226, 227, 235, 278, 292, 294, 611,
614, 617; Bed II of, 227, 278,
610, 611, 613, 614, 617; Bed V
of, 278, 634, 635, 636; Capsian
skeleton from, 278, 634, 635, 636;
Chellian-3 skull from, 336, 337,
614, 616-17; dating of deposits in,
314; milk teeth from, 610, 613
Oligocene epoch, 32, 189, 190, 191,
193, 194. 196
Omo fauna, 225, 228, 234, 235, 295
Onges, 425
ontogeny, phylogeny recapitulated
by, 164-5
Oppenoorth, W. F. F., 392, 398
orangs, 140, 142, 143-4; ancestors
of, 198 ff.
Ordos sites, 470-1
Oreopithecus, 209-15, 334, 380
Oriental faunal region, 50, 52, 53,
54, 56, 57, 92, 121, 136, 143, 189,
211,373, 421,485
Origin of Species, 50
Orochons, 65
osteometry, standardized by Martin,
516
Ostiaks, 451
overlap test, for subspecies, 16-17
Paidopithex, 203
Pakistan, 2, 138, 422, 423, 482, 484
Paleanthropic grade, 334, 336
Palearctic faunal region, 50, 52, 53,
54, 56, 57, 149, 221, 485
Paleocene epoch, 32, 189, 190, 191
Paleolithic tools, 324-32, 488
Paleosima, 203
Paleozoic era, 187 n.
Palestine, 222, 229, 301, 304, 330,
333, 484, 488, 526, 555, 577, 578,
604; inhabitants of, during
Wiirm I, 13, 103, 565-75, 576;
Upper Paleolithic fossil man sites
in, 581, 587
Panama, Isthmus of, 43, 190
Panganrejang Cave, 415
Papio, 137, 139-40, 195
Papuans, 2, 44, 112, 367, 369, 407,
420, 425, 426, 432
parallel evolution, 11, 37, 132, 192
Parapithecus, 193, 196
parasites, mutual, on primates, 176,
193
patas monkey, 137
Pederson, P. O., 357
Pedicinus, 176
Pediculus, 176
pedomorphism, 25, 647
Pei, W. C., 207, 239, 431, 465, 467
Penck, Albrecht, 310, 311, 312, 314
Perigordian industry, 579
peripheral gene flow, 36, 37
Persian Gulf, 54
pharynx, 74, 75
Philippines, 52, 112, 139, 230, 415,
425
Pho Binh Gia skulls, 420
Phthirus, 176
phyletic evolution (succession), 27,
28, 106 ff., 111
phylogeny, recapitulated by ontog-
eny, 164-5
physiological adaptation, to altitude,
70-1; to cold, 59-68, 69; to heat,
68, 69
phytography, 4 1 n.
Pilling, A. R„ 97
Pinjor fauna, 222, 223
Pithecanthropus, 10, 13, 323, 336,
337; cranial capacity of, 393; sites
of skeletal material (table), 376;
Index
xv
Pithecanthropus ( continued )
teeth of, 344, 387-90, 453, 454,
455; thighbones of, 386
Pithecanthropus B, 375, 380-2, 388,
389, 449, 450, 453, 454
Pithecanthropus 1, 384, 385, 386
Pithecanthropus 2, 284, 384, 385
Pithecanthropus 3, 384
Pithecanthropus 4, 374, 375-80, 384,
388, 453
Pithecanthropus dubius, 382, 387,
388, 389
Pithecanthropus erectus, 10, 384
Pithecanthropus modjokertensis, 299,
383-4; see also Homo modjoker-
tensis
Pithecanthropus pekinensis, 437
Pithecinae, 133
Pitjendjera tribe, 66, 67
Piveteau, J., 410, 437
platyrrhines, 131-3; evolution of,
192
Pleistocene epoch, 4, 32, 33, 34, 42,
44, 5°, 52, 55, 56, 189, 207, 218,
313; and apes of China, 206-7;
baboons in, 195; close of, 579,
590; first glacial advances of, 52,
479; human evolution during, 57,
78, 101; hunting begun, 80;
Lower, see Lower Pleistocene
epoch; macaques in, 195; Meso-
pithecus in, ig5; Middle, see Mid-
dle Pleistocene epoch; migrations
of early man during, 58; pluvial
periods in Africa during, 315; Up-
per, see Upper Pleistocene epoch
Pliocene epoch, 11, 32, 47, 189, 190,
205, 218, 221, 222, 225, 283, 310;
baboons in, 195; Dryopithecus in,
202, 203; Fort Ternan primate in,
205-6; and Italian fossil beds,
209; Java submerged during, 223;
Lower, 204, 205, 209, 219; ma-
caques in, 195; marsupials in, 92;
Mesopithecus in, 195; Ramapithe-
cus in, 204, 219; Upper, 218, 219
Pliopithecus , 197, 198
pollen analysis, of Choukoutien brec-
cia, 436; of Florisbad deposits,
644
polymorphism, balanced, 22-3; de-
fined, 14 n.
Polynesians, 2, 62, 353, 355, 367,
369
polytypic species, 14
Pongidae, 140, 141
Ponginae, ig9
Pongo pygmaeus (orang), 140, 142,
143-4
Poppy, A. J., 634
population, in taxonomy, 9
porpoise, brain of, 78-9
Portugal, Neanderthals in, 527
postnatal growth, differences in,
171-2
posture: erect, see erect posture;
summary of data on, 154-62
precipitin fest, 174, 193
premolar cone, 356-7
primates, 119-50; behavior of, as
criterion of species, 183-5; chro-
mosomes studied in, as new tool
of taxonomy, 177-83; classification
of, 120 122; fossil record of,
from lemurs to swamp apes, 186-
216; parasites on, mutual, 176,
3.93; physiological tests on rela-
tionships among, 172-6; man’s
place among, 151-85; sexual be-
havior of, 82-4, 183, 184; sexual
selection among, 85-6
Proconsul, 199-202, 216-19 passim,
248, 252, 254, 262, 264, 268,
285, 290
Propliopithecus, 196
prosimians, 120-1; proliferation of,
190-2
Proterozoic era, 187 n.
Protoanthropic grade, 334, 336
Ptilocerus, 121
puberty ceremony, 89
Punans, 421
purine metabolism, 172, 193
Putjangan beds, 316
Pycraft, W. P., 626
Pygmies, 4, 5, 13, 26, 34, 588, 589,
651-5, 658; achondroplastic, 115,
653; ateliotic, 115; birth rate of,
652; classification of, 112; distri-
bution of, in refugee pattern, 57;
cranial capacity of, 654; faces of,
654; as food gatherers, 99; hair
of, 654; and Negroes, gene flow
between, 652, 653; number of,
651, 660; pelvic bones of, 242-3;
XVI
Index
Pygmies ( continued )
sexual dimorphism in, 653; sickle-
cell found among, 652; skin color
of, 654; stature of, 653
Pyrenees, 527
quadrupedal primates, descent from,
153
Quaternary period, 188 n.
Queensland, 408
Rabat remains, 596-8
racial differences, in living men, 662
racial types, defined, 19
rain forest, 44, 52, 589, 630
Ramapithecus, 203-5, 2°6, 215, 219,
223
Ratcliffe, H. L., 109
recapitulation law, propounded by
Haeckel, 164-5
Recent epoch, 32, 33, 189, 218, 305,
476-7, 630
Reck, Hans, 634
recombination, defined, 21
Remane, A., 207
Rensch, B., 57
Rhinopithecus, 136
Rhodesian man, see Broken Hill man
Riffians, 601
Rift Valley, 315
Riss glaciation, 309-16 passim, 318,
320, 477, 478, 480, 486, 527
Riss-Wiirm (Last) Interglacial, 310,
3H, 318, 323, 329, 390, 460, 486,
497, 498, 5°i, 505, 507, 508,
521, 522, 523; mandibles of Eu-
ropeans living in, 511-14; and
postcranial bones from Krapina,
516-19; teeth of Europeans living
in, 514-16
Riss-Wiirm Interstadial, 497
rites of passage, 89-90
Robinson, J. T., 231, 240, 241, 243,
244, 252, 267, 270, 271, 295, 301
Roche, Jean, 595
Rocky Mountain system, 189
Roginskii, 1. 1., 558
Ruanda-Urundi, 651
Rumania, 525, 553, 581
Rusinga Island, 196, 199
Saale glaciation, 222 n., 310
Saccopastore remains, 487, 500-4,
514, 515, 522, 527
Sahara, 43, 52, 320, 321, 589, 590,
607, 635, 636, 637, 649, 660
Sahul Shelf, 44, 46, 56, 92, 190,
318, 399, 406
Sakai, 469
Salawati, 55
Saldanha Bay skull, 337, 619-21
Sampoeng F cranium, 414
Sandawe tribe, 4, 648
Sangi Island, 55
Sangiran, 299, 375, 382, 389
Sangoan industry, 330
Santal tribe, 420, 422
sapiens-erectus threshold, 16, 337-
46, 427, 633, 657; and brain size,
337_4i; and cranial form, 341-4;
and tooth size, 344—6
Sarasin, Paul and Fritz, 414
Schepers, G. W. H., 238
Schlosser, M., 196
Scholander, P. F., 64, 66
Schultz, A. H., 145, 151, 166, 167,
170, 212, 214, 215
Schwalbe, G., 214
Schwidetzky, Ilse, 111 n.
Sclater, P. L., 50
Second Himalayan Glaciation, 229,
310
Second Interglacial, see Mindel-
Riss Interglacial
selection, defined, 21-2
Semang, 112
§enyiirek, M. S., 295, 561
Sergi, Sergio, 334 n., 501, 502, 504
serum albumin and serum gamma
globulin tests, 174
serum transferrins, in primates, 175
Sewall Wright effect (genetic drift),
47-8
sexual dimorphism, 26-7
sexual selection, 85-6
Shanidar, Eastern Neanderthals of,
103, 561-5, 576
Shibar pass, 53
shoveled teeth, 355-6
shrinking process: in Capoids, 645;
in Pygmies, 654
Shukba remains, 567
Siam, 52, 53, 143, 230, 330, 420
siamangs, 140, 141, 142, 143
Index
XVII
Siberia, 54, 318, 330, 331, 577
sickling trait, 22, 70, 652
Sidi Abd er-Rahman, 344
Sikkim, 18
Simons, E. L., 204, 205
Simpson, G. G., 10, 120, 193, 198,
203, 437
Sinanthropus, 10, 13, 85, 207, 317,
336, 337> 340. 382, 429; brain
case of, 438-45; face of, 445-7;
features in common with living
Mongoloids, 458-60; fire possessed
by, 91, 436; humerus of, 457; leg
bones of, 456-7; longevity of, 103,
mandibles of, 447-52; position of,
in human family tree, 458—60; tax-
onomy of, 437-8; teeth of, 344,
452-6, 516
Sinanthropus pekinensis, 10, 430-6,
437; age of specimens of, 433;
loci of specimens of, 434; sex de-
termination of, 433, 434
Sinelnikov, N. A., 561
Singa skull, 639-40
Singer, Ronald, 393, 619, 620, 630,
633, 645
Singhalese, 423, 424
Sino-Malayan fauna, 223
Sipka Cave, 550, 552
Sivapithecus, 203; africanus, 219,
293
Siwalik Hills, India, 195, 203, 218,
222, 223
Sjara-Osso-Gol tooth, 470-1
Skerlj, B., 508
Skhul postcranial skeletons, 572-5,
576
Skhul skulls, 569-71
skull, defined, 256
slow game, defined, 80
Smith, Eliot, 623
Smugglers’ Cave mandible, 595-6
Snow, Charles, 610
social adaptation, evolution through,
72-118 passim
social structure: energy converted
into, 90; speech related to, 88
soil analysis, 234, 235
Solecki, Ralph S., 562
Solo leg bones, 397-8
Solo River Valley, 315, 390
Solo skulls, 336, 337, 340, 390-9;
age of, 391, 392; cranial capacity
Solo skulls ( continued )
of, 393; faces removed from,
396-7; hypophyseal fossa of, 395;
internal dimensions of, 439; in-
juries to, 391-2; racial anatomy of,
392-6; sex of, 391, 392, 393
Solutrean industry, 579
South Africa, 217, 226, 228, 589,
590, 631, 653, 660; Capoid skele-
tal material in, 637, 645, 647;
climate of, 52, 315; Levalloisio-
Mousterian industry in, 330; re-
moteness and isolation of, 630
South America, 54, 477, 478, 480;
area of, 43; dwarf marmoset in,
113; tool industries in, 331, 478;
land mass of, 42, 43, 46; monkeys
of, 131-3; North America con-
nected to, 43, 190
South Asia: dwarf populations in,
34; and exchange of animals with
Africa, 56; Palearctic genera in,
57; spiral-haired peoples of, 4
Southeast Asia, 20, 44, 56, 112, 121,
330, 373, 421, 425
Southern Hemisphere, 46, 57, 68, 92,
311, 410
Soviet Union Neanderthal remains,
525, 554-61, 576; in Kiik-Koba
cave, 555-7; in Starosel’e, 557-8;
in Teshik-Tash cave, 558-61, 576
Soviet Union Upper Paleolithic sites,
581
space dimension, in study of human
origins, 318-22
Spahni, J-C., 549
Spain: Neanderthals in, 525, 526,
527; Upper Paleolithic fossil man
sites in, 580
special adaptation, 28
species, allopatric, 14; differentiation
of, 21-2; euryphagous, 15; geo-
graphical differentiation of, 14,
15; interbreeding of, 12-13; inter-
fertile, 12; life spans of mamma-
lian, 32-4; monotypic, 14; poly-
typic, 14; spatial requirements of,
14, 15; stenophagous, 15; sym-
patric, 15; in taxonomy, concept
of, 9, 11-13
speech: hunting related to, 80, 87;
invention of, 73, 74, 76, 80, 87;
Index
xviii
speech ( continued )
learning of, 73-4; organs of, 74-5,
76; and puberty ceremony, 89;
social structure related to, 88; see
also language (s)
spider monkeys, 132, 133
Starosel’e, infant skeleton of, 557-8
Steinheim cranium, 341, 487, 492-5
stenophagous species, 15
step flaking, 501
Sterkfontein site, 232, 233, 234, 235,
236, 238, 239; capitate bone from,
253; femoral bones from, 245,
246; humerus from, 251; pelvic
bones from, 241, 243; scapula
from, 249, 250; skulls from, 257,
258; teeth from, 267, 270, 271;
vertebrae and ribs from, 240-1
Stewart, T. D., 480, 563, 564, 565
Stillway industry, 622, 628, 631,
632
Stirling, Matthew, 278
Strandloopers, 637, 646
Straus, W. L., Jr., 151, 227
stress, study of, 552-3
Subalyuk cave: child’s skeleton
from, 552-3; mandible from, 551;
postcranial bones from, 552
subspecies: concept of, 9, 15-17;
successional, 17
sub terminal chromosome, 178
succession, evolutionary mechan-
isms of, 27, 28, 106 ff-, 111
successional subspecies, 17
Sudan, 320, 639, 650
Suez, Isthmus of, 42, 43, 190
Sugrivapithecus, 203, 346
Sumatra, 52, 142, 143, 330, 415, 421
Sunda Shelf, 44, 46, 56, 92, 143,
190, 318, 406
Suzuki, H., 464, 472, 476, 566
Swanscombe skull, 91, 314, 487,
495-7
Swartkrans site, 232, 233, 234, 235,
236; metacarpal of thumb from,
253, 254; pelvic bones from, 241,
243; skulls from, 267, 270, 271
Switzerland, 525, 580
symbiosis, 100
sympatric species, 15
Symphalangus, 142
Syria, 330, 484, 505, 577, 578
Tabun material, 566, 567, 568-9,
57i
Taforalt remains, 600, 605, 607, 600
Takai, F., 464
talapoin, 137
Talaud Island, 55
Talgai skull, 408-9, 410
Tam Hang site, 417-19
Tam Pong skull, 417
Tamils, 354, 423
Tanganyika: Olduvai Gorge in, see
Olduvai Gorge; Sandawe tribe in,
4, 648
Tangier man, 598-600, 601, 602
tarsiers, 120, 129-30
Tartangan culture, 406
Tasmania, 31, 44, 55
Tasmanians, 4, 31, 34, 112, 306,
425; and facial flatness, 367; geo-
graphical distribution of, 59;
woolly hair of, 44
Tatrot fauna, 222, 223
Taubach tooth, 514
Taung site, 232, 233, 234, 235, 236,
238, 239; skull from, 257, 258,
263; teeth from, 270
taurodontism, 359-60, 455, 456, 516
taxonomy, of Australoid subspecies,
425-7; and behavior study of pri-
mates, 183—5; and chromosome
study of primates, 177-83; prob-
lems of, 9—10; of Sinanthropus,
437-8; single-character, obsolete
concept of, 13; species concept in,
9. n-13
Tayacian remains, 499, 500, 501,
521
Tayaki, F., 472
Tchad, Australopithecine from, 297,
334> 590
teeth: of Ainu, 355, 357, 516; of
Australian aborigines, 344, 359,
362, 363; of Australoids, 352, 353,
426, 453; of Australopithecines,
256, 267-77, 352, 357, 359, 360;
bearing of, on erect posture,
153-4, 162-4; °f Broken Hill man,
624-5; of Bushmen, 344, 353, 354,
359, 360, 362, 364, 455, 456; of
Capoids, 354, 360, 364; of Cau-
casoids, 352, 353, 354, 355, 360,
361, 362, 363, 364; cynodont,
359; of Dryopithecus, 203, 219,
Index
xix
teeth ( continued )
360; of Eskimo, 355, 357, 360,
362, 363, 455, 456, 516; of Euro-
peans of Last Interglacial age,
514-16; of Fort Ternan primate,
205- 6; of gibbons, ancestral, 196,
197; of Gigantopithecus blacki,
206- 7; Haua Fteah, 603; of In-
dians, American, 369, 474; mega-
dont, 353; of Meganthropus, 298-
300; of Melanesians, 353; meso-
dont, 353; of Mesopithecus, 195;
microdont, 353; of Mongoloids,
352, 353, 355, 356, 357, 359, 360,
362, 363, 364, 428, 453; morpho-
logical differences in, 350-64 pas-
sim; Mount Carmel, 571-2; of
Neanderthals, 455, 456, 539-42;
of Negritos, 353; of Negroes,
353-4, 356, 360, 361, 362, 363;
from Olduvai Bed II, 610—11, 613;
of Olduvai child, 282-3; of Oreo-
pithecus, 211-12; of Pithecanthro-
pus, 344, 387-90, 453, 454, 455;
of Polynesians, 353, 355; of Pro-
consul , 201; of Rabat man, 596-8;
racial variations in form and struc-
ture of, 354-64; of Ramapithecus,
204-5, 219; shoveled, 355-6; of
Sinanthropus, 344, 452-6, 516;
size of, and sapiens-erectus thresh-
old, 344-6; of Sivapithecus africa-
nus, 219; of Tangier man, 598-
600; taurodont, 359-60, 455, 456,
516; Ternefine, 593-4; Ting-tsun,
460-1; of Upper Paleolithic Euro-
peans, 584; of Wadjak man,
404-5; of Zinjanthropus, 292-4,
352, 359
Telanthropus, 233, 252, 264, 267,
270, 282, 289, 299, 323
Tell Ubeidiya, fossil hominid of,
229, 237, 297-8
telocentric chromosome, 178, 179
temperament, and endocrines, 115-
16
ten Haar, C., 390, 392
Ternefine remains, 344, 441, 452,
489, 5!3, 591_5, 600, 601, 602
Ternefine-Tangier line, 600-2, 649,
658
Terra, M. de, 212
Terra Rossa, 317
Tertiary period, 32, 44, 54, 56, 57,
188 n., 191
Teshik-Tash cave, Neanderthal boy
from, 558-61, 576
Thenius, E., 214
Theropithecus, 137, 140
Thieme, F. P., 35
Third Interglacial, see Riss-Wiirm
Interglacial
thumb, 159, 160; in fetal life, 167
Tian Shan Mountains, 43, 54, 482,
554
Tiberias, Lake, 79-80, 229, 566
Tibet, 44, 70, 207, 208, 316, 318;
hoolock in, 143; land mass of, 43;
langur in, 136
Tibetans, 26-7, 54, 70, 661
Tierra del Fuego, 64, 69, 428
time dimension, in study of human
origins, 309-18
Timor, 56, 113, 406
Ting-tsun man, 317, 460-1, 521
Ti-Shao-Gou-Wan remains, 471
Tiwi society, 94-9, 100, 102, 184,
411
Toala, 414, 415
Tobias, P. V., 296, 641
tool association, 234, 235
tool-making, 76, 77, 90, 227, 306,
307-8, 327-32; by Australopithe-
cines, 237-g; in Lower Pleisto-
cene, 227-30, 333; paleolithic,
324-32, 488
torus mandibularis, 451, 452, 512,
537
Tratman, E. K., 357
Tratz, E., 146
tree shrews, 121, 126
Trevor, J. C., 603
Triassic period, 188 n.
Trinidad, 54
Trinil fauna, 224, 260, 314, 316,
337, 384-6, 387 ff-
tubera frontalia, 509-10
Tunisia, 320, 522, 607
Tupaia, 121
Turkey, 482, 484, 526, 562, 577, 578
Turkomans, 18
Twiesselmann, F., 654
Tze-Yang skull, 337, 465-7
XX
Index
Upper Cave of Choukoutien, 337,
472-5
Upper Paleolithic Europeans, 347,
488, 577-87; as artists, 585—6;
Asiatic relatives of, 587; descend-
ants of, 587; fossil sites of, 580-2;
height of, 582-3; racial character-
istics of, 582-6; skeletons of,
583-4
Upper Pleistocene epoch, 305, 316,
317, 320, 321, 323, 328, 329, 390,
412, 484, 630, 660; Choukoutien
Upper Cave people from, 472-5;
European fossil men of, 497 ff.;
Honshu remains from, 471-2; Liu-
Kiang man from, 467-70; Niah
cave skull from, 412, 413; and
Ordos sites, 470-1; Tze-Yang
woman from, 465-7
Urey, Harold C., 312
Urogale, 121
Ushikawa quarry, humerus shaft
from, 464-5
Uzbekistan, 484, 558, 559
Uzbeks, 18, 144, 228, 651
Vallois, H. V., 260, 410, 437, 500,
512, 517, 523, 607, 608, 628, 650
variety, as term used in taxonomy, 9
Veddas, 99, 366, 414, 423, 424, 518
Venezuela, 478
vertebral column, and posture,
154-5, 156, 157, 159
Villafranchian fauna, 207, 221, 222,
229, 297, 316, 321
Vlcek, E., 507
von Bonin, G., 582
von Koenigswald, G. H. R., 223,
226, 227, 230 n., 299, 300, 380,
382, 392, 398, 610
Waagen, W., 17
waagenons, 17, 334
Wadjak man, 399-406, 407, 408,
413; brain case of, 401-3; denti-
tion of, 404-5; face of, 403, 427,
445; mandible of, 403-4, 449; sig-
nificance of, 405-6
Waigeo, 55
Wallace, A. R., 50, 55
Wallacea, 55-6, 92, 401, 406, 522
Wallace’s Line, 55, 373
Washburn, S. L., 80, 139, 140, 151,
235, 243, 244 n., 258, 260, 395,
437
Watusi, 13, 636
Weber’s Line, 55
Weichsel glaciation, 310
Weidenreich, Franz, 378, 382, 389,
391-5 passim, 400, 402, 407, 410,
429-35 passim, 447-8, 452, 456,
460, 480, 506, 627; quoted, 398,
437, 440 n., 474-5
Weinert, H., 295, 493, 627
Wells, L. H., 603
West Africa, 589, 590; monkeys in,
135, 140; Negroes in, 22, 590,
633; Sangoan industry in, 330
West Indies, 54
West Pakistan, 2, 423, 482, 484
Western Asia, as Caucasoid territory,
482, 484; fossil men in, 487, 498,
587; as nuclear region, 485
Western European Neanderthals,
527-49; build of, 548; clavicles of,
543; crania of, 529-35; extinction
°f, 548-9; faces of, 534-5; feet of,
546-7; femora of, 546; hands of,
545; height of, 548; humeri of,
544; kneecaps of, 546; mandibles
°f, 535-9; noses of, 533-4; pelvic
bones of, 545-6; postcranial skele-
tons of, 542-7; radii of, 544; ribs
of, 543; teeth of, 539-42; tibia of,
546; ulnae of, 544; wrist bones of,
545
whole globulin molecules, in pri-
mates, 175
Wicker, Fred, 205
Willey’s Kopje, 636
Wilton culture, 330, 631, 638, 640,
641
Wisconsin glaciation, 310, 312, 313,
320, 477, 479
Wolof skulls, 650
Woo, Ju-Kang, 203, 449, 463, 464,
467, 468, 469, 476
Woo, T. L., 365
Wood-Jones, 151
Wormington, H. M., 479
Wiirm glaciation, 68, 309-15 passim,
318, 320, 476-9 passim, 486
520-3 passim, 526, 548, 550, 579,
Index
xxi
Wiirm glaciation ( continued )
582; inhabitants of Palestine dur-
ing. 565-75
Yakkhas, 423
Yellow Earth, Age of, 317
Yerkes, Robert M., 146
Yeti, 208
Yugoslavia: Krapina remains from,
487, 508-11, 513-14, 515, 516-
19; Veternica skulls from, 554
Zagreb Museum, 508, 510
Zagros Mountains, 526, 562, 577
Zapfe, H., 198
Zeuner, F., 311, 312, 313
Zinjanthropus, 80, 226, 278, 281,
287-9. 385. 617; cranium of, 289-
92; diet of, 288; face of, 289; leg
bones attributed to, 294; status of,
294; teeth of, 292-4, 352, 359,
389; tools of, 288
Zitzikama collection, 645
zoogeography, 41, 49, 58
A NOTE ABOUT THE AUTHOR
Carleton Stevens Coon, curator of ethnology and professor of
anthropology at the University Museum in Philadelphia since
1948, was born in Wakefield, Massachusetts, and received his
A.B., A.M., and Ph.D. from Harvard University. He has divided
his time between field work and teaching, first at Harvard, then
at the University of Pennsylvania. In connection with his work
he has traveled extensively in Africa, Asia, and Europe. Dr.
Coon arranged the famous Hall of Man exhibit at the University
Museum, and is a regular panel member on the Peabody
Award-winning television program What in the World? He is
President of the American Association of Physical Anthropolo-
gists. Among his many books are Caravan: The Story of the Mid-
dle East ( i951, 1958); The Story of Man ( 1954, revised edition
1962); and The Seven Caves (1957). Dr. Coon has been a
leading authority on race ever since his famous and highly suc-
cessful book, The Races of Europe, was published in 1939. He
has two sons and five grandchildren, and lives with his wife,
the fonner Lisa Dougherty, in West Gloucester, Massachusetts.
September ig6 2
A NOTE ON THE TYPE
The text of this book is set in Caledonia, a Linotype face
designed by W. A. Dwiggins (1880-1956), the man respon-
sible for so much that is good in contemporary book design and
typography. Caledonia belongs to the family of printing types
called “modern face” by printers— a term used to mark the
change in style of type-letters that occurred about 1800. Cale-
donia borders on the general design of Scotch Modem but is
more freely drawn than that letter.
Composed, printed, and bound by
Kingsport Press, Inc., Kingsport, Tennessee.
Typography and binding design
based on originals by
W. A. DWIGGINS
(continued from front flap)
many kinds of cognate evidence, the parallelism
of the evolution of the separate races soon be-
came apparent. And the prehuman relatives of
man were drawn forward in time to the very
date of the earliest human skull.
In the process of proving his general theory,
Dr. Coon has also produced the first detailed
history of the evolution of the five races of man.
This book, then, is not alone a vast scientific
synthesis, but a work of history: the history of
a primate genus. And in it science serves only
as a set of tools for reconstructing the pathways
of human evolution.
CARLETON STEVENS COON
Curator of Ethnology and Professor of Anthro-
pology at the University Museum in Philadel-
phia since 1948, was born in Wakefield, Massa-
chusetts. He obtained his A.B., A.M., and Ph.D.
from Harvard University, where he also taught
before he went on to the University of Penn-
sylvania. Dividing his time between teaching
and field work, he has traveled extensively in
Africa, Asia, and Europe. He arranged the
famous Hall of Man exhibit at the University
Museum, and is a regular panel member on the
Peabody -Award -winning television program,
What in the World ? He has served as President
of the American Association of Physical An-
thropologists, and is a leading authority on
race. Among his books are Caravan: The Story
of the Middle East (1951, 1958); The Story of
Man (1954, Second Edition, 1962); and The
Seven Caves (1957). k>r- Coon has two sons and
five grandchildren, and lives with his wife, the
former Lisa Dougherty, in Devon, Pennsylvania.
Alfred A. Ktiopf, Publisher
NEW YORK
PRINTED IN U.S.A.
FROM THE FIRST CHAPTER OF
THE ORIGIN OF RACES
At the dawn of history, which is another way of saying “beginning
with Herotodus,” literate people of the ancient world were well aware
that mankind was divided into a number of clearly differentiated
races. Even before that, racial differentiation can be traced back to
at least 3000 b.c., as evidenced in Egyptian records, particularly the
artistic representations. We also have pictures of white people on the
walls of western European caves which are as much as 20,000 years
older.
How many kinds of people there were in the world was not
really known until after the voyages of discovery which tore the veil
from the Americas, the Pacific Islands, and Australia. Even then, the
problem of classifying the races remained, and it has not been settled
to this day.
For present purposes I am using a conservative and tentative
classification of the living peoples of the world into five basically geo-
graphical groups: Caucasoid, Mongoloid, Australoid, Congoid, and
Capoid. The first includes Europeans and their overseas kinsmen, the
Middle Eastern whites from Morocco to West Pakistan, and most of
the peoples of India as well as the Ainu of Japan. The second includes
most of the East Asiatics, Indonesians, Polynesians, Micronesians,
American Indians, and Eskimo. In the third category fall the Austral-
ian aborigines, the Polynesians, Papuans, some of the tribal folk of
India, and the various Negritos of south Asia and Oceania. The fourth
comprises the Negroes and Pygmies of Africa. The fifth group includes
the Bushmen and Hottentots and other relict tribes like the Sandawe
of Tanganyika. . . .
My aim in this book is to see how far back in prehistoric an-
tiquity human racial types can be traced. Did they all branch off a
common stem recently ... or did their moment of separation lie lower
down on the time scale when long-extinct types like the so-called ape-
men of Java and China were still alive?
All of the evidence available from comparative ethnology, lin-
guistics, and prehistoric archaeology indicates a long separation of the
principal races of man. This is contrary to the current idea that the
archaic species of man who had preceded Homo sapiens became con-
veniently extinct.
Man is little more than a half million years old. Geologically
speaking, we were bom yesterday. Fossil men now extinct differed
from each other in race and were not members of separate species,
except in the sense that one species grew out of another.
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